Guancha, MIKLUCHO-MACLAY, 1868

Rapp, Hans Tore, 2006, Calcareous sponges of the genera Clathrina and Guancha (Calcinea, Calcarea, Porifera) of Norway (north-east Atlantic) with the description of five new species, Zoological Journal of the Linnean Society 147 (3), pp. 331-365 : 349-358

GUANCHA MIKLUCHO- MACLAY, 1868

Type species: Guancha blanca Miklucho-Maclay, 1868 . Clathrinidae with a cormus composed of a peduncle and a clathroid body. The peduncle may be formed by true tubes with a normal choanoderm, or may be solid with a special skeleton. The skeleton is composed of regular (equiangular and equiradiate) spicules to which parasagittal spicules may be added, at least in the peduncle. In some species, only parasagittal spicules are present. The unpaired actine of parasagittal spicules is always basipetally orientated.

Remarks on Guancha

Prior to this work, the genus was represented by ‘about six’ species ( Borojevic et al., 2002). Species within the genus Guancha display a progressive dif- ferentiation into a clathroid body and a peduncle. In the more simple species, such as G. arnesenae sp. nov., G. blanca Miklucho-Maclay, 1868 and G. sagittaria ( Haeckel, 1872) , the peduncle is composed of normal tubes with a complete choanoderm. G. blanca is only slightly different from a typical Clathrina , and only the fact that it represents the starting point for an evolutionary line makes it possible to separate it into a distinct genus ( Borojevic & Peixinho, 1976). In more morphologically complex species such as Guancha challengeri ( Poléjaeff, 1883) , G. lacunosa ( Johnston, 1842) and G. pulcherrima (Dendy, 1891) the peduncle acquires a solid form, consequently with no choanoderm, and with a particular skeleton. Guancha is also characterized by possessing parasagittal triactines organized in a parallel manner with the unpaired actine basipetally orientated. In G. blanca the parasagittal spicules are found in the peduncle only, whereas in species such as G. arnesenae sp. nov., G. tetela Borojevic & Peixinho, 1976 and G. sagittaria all the spicules are parasagittal.

The classification of Guancha is especially difficult when the skeleton is composed of more-or-less uniform spicules with only slightly longer unpaired actine in the peduncular triactines. G. blanca has been reported from most oceans, from the littoral zone to the abyssal plains ( Barthel & Tendal, 1993; Janussen, Rapp & Tendal, 2003). Cosmopolitanism is very rare among sponges ( Klautau et al., 1999), and in the closely related Clathrina the use of statistical analyses of shape, size and distribution of spicules, combined with molecular methods, have shed new light on the classification of the genus, and have revealed that geographical distribution of the different species is more restricted ( Klautau et al., 1994; Wörheide & Hooper, 1999; Rapp et al., 2001; Klautau & Valentine, 2003). In agreement with Borojevic et al. (2002), I consider that G. ‘ blanca ’ represents a complex of species, and that the ongoing work revising the genus, including analysing specimens of G. ‘ blanca ’ from different geographical regions and from shallow to abyssal depths with a morphological and molecular approach, will reveal a number of new species.

GUANCHA ARNESENAE SP. NOV.

( FIGS 13A, B , 14 , TABLE 7)

Type locality: Evenskjer, 68°35′N, 16°35′E, 25 m.

Holotype: VM-398 , Evenskjer, 68°35′N, 16°35′E, 25 m, 18.08.1919, 1 specimen.

Paratype: TM-183, Ramfjorden, 10–20 m, 14.07.1921, 1 specimen.

Etymology: Named in memory of the Norwegian spongiologist Emily Arnesen (1867–1928).

Previous records: None.

Additional material examined: 28 specimens.

IB. Evenskjer, 68°35′N, 16°35′E, 25 m, 18.08.1919, 2 specimens . TMU. TM-150, Ramfjorden, 69°33′N, 10°10′E, 10–20 m, 14.07.1921, c. 10 specimens (small and fragmentary) . VM. VM-16904 , Tromsø, 69°40′N, 19°00′E, 30–35 m, 28.10.1916, 1 specimen . VM-16905 , Tromsø, 69°40′N, 19°00′E, 30 m, 21.09.1917, 1 specimen . VM-16906 , Gibostad, 69°21′N, 18°05′E, 50 m, 23.05.1912, 4 specimens (dry) . VM-16907 , Lødingen, 68°24′N, 15°58′E, 70–90 m, 22.08.1913, 1 specimen (dry) . ZMO. B-1289, Balsnes, 69°34′N, 18°52′E, 40– 60 m, 9 specimens .

Description

Clathrinidae with a cormus normally composed of a clathroid cormus of irregularly and loosely anastomosing tubes and a peduncle. Size ranging from 4 to 10 mm in height. Several true tubes with a normal choanoderm form the peduncle. The peduncular tubes run in parallel, sometimes free, or slightly anastomosed. In specimens without a proper peduncle the cormus is narrower at the base than in the apical region. Water-collecting tubes converge at the centre of the sponge, ending in one or several apical oscula. The single tubes are about 0.5 mm in diameter. Colour is light beige in alcohol and when dried. Texture soft. The skeleton of the cormus is solely composed of tripodic, parasagittal triactines. The spicules are irregularly orientated in several layers in the walls of the tubes, resulting in a wall thickness of about 30–50 µm. In the peduncle the skeleton consists of tripodic parasagittal triactines, generally similar to those in the cormus, but with shorter paired actines. In addition, there are very characteristic parasagittal triactines where the paired actines are bent, forming a concave ‘U’. Spicules in the peduncle with their unpaired actine basipetally orientated. All the actines are straight, cylindrical and with blunt tips.

Distribution

The species is found in the northern part of Norway at 10–90 m depth.

The species is also known from Greenland at 30– 54 m depth (my unpubl. data).

Remarks

The species is very similar to Guancha sagittaria Haeckel, 1872 . However, there are some differences. Haeckel (1872) made his original description based on three very small olynthus tubes, dredged from Storebaelt ( Denmark) during the Pommerania- Expedition in 1871, probably representing very young specimens. All the specimens examined here have clathroid cormi, and a more-or-less developed peduncle. Both species have the typical shape of the parasagittal triactines from the peduncle, but whereas all spicules are cylindrical in G. arenesenae they are conical in G. sagittaria ( Haeckel, 1872) . The presence of only parasagittal triactines in G. arnesenae differentiates it from all other species of Guancha except Guancha tetela Borojevic & Peixinho, 1976 from Brazil. However, G. arnesenae and G. tetela are easily distinguished by the shape of the paired actines of the parasagittal triactines from the peduncle, forming a concave ‘U’ in arnesenae and close to a ‘T’ in tetela .

GUANCHA BLANCA MIKLUCHO- MACLAY, 1868

( FIGS 15A–D , 16 , TABLE 8)

Original description: Guancha blanca Miklucho- Maclay, 1868: 221–232, plates 4 and 5.

Synonyms and citations

Ascetta blanca ( Hansen, 1885: 20; Arnesen, 1901a: 9–10).

Clathrina blanca ( Ereskowsky, 1995: 730) .

Guancha blanca ( Borojevic & Boury-Esnault, 1987: 14–15; Borojevic et al., 1990: 252; Barthel & Tendal, 1993: 84; Rapp, 1999: 65–71; van Soest, 2001: 101; Borojevic et al., 2002: 1143–1144; Janussen et al., 2003: 17–19).

Leucosolenia blanca ( Levinsen, 1886; Breitfuss, 1898a: 105–106, 1898b: 13–14, 1898c: 295–296, 1911: 224; Derjugin, 1915: 289; Breitfuss, 1927: 27, 1933: 240, 1936: 5; Alander in Jägerskiöld, 1971: 60).

Leucosolenia lacunosa ( Burton, 1930b: 488) .

Leucosolenia macleayi ( Burton, 1930a: 14–17, 1930b: 487; Arndt, 1935: 8–9).

Type locality: Lanzarote, Canary Islands.

Holotype: The type is probably lost (A. Plotkin, ZIL, pers. comm.).

Previous records: Bergen and Stavanger ( Burton, 1930b; as L. macleayi ); Singlefjord ( Breitfuss, 1936).

Additional material examined: 147 specimens.

IB. Skarnsundet, 63°53′N, 11°04′E, 200 m, 09.10.1968, 3 specimens . Trondheimsfjord, c. 63°30′N, 10°30′E, 35 different localities, 14–500 m, 1995–96, 126 specimens. Nord-Leksa, 63°35.9′N, 09°26.3′E, 150–200 m, 21.03.1996, 2 specimens. Korsfjord, 60°09.12′N, 05°08.52′E, 300 m, 12.02.2003, 1 specimen. Balsnes, 69°34.1′N, 18°52′E, 40–60 m, 1 specimen. TMU. TM-180, Beiarn, 67°04′N, 14°35′E, 35–45 m, 02.08.1955, 2 specimens . TM-181, Trondenes, Vågsfjord, 68°50′N, 16°35′E, 90–150 m, 28.08.1954, 2 specimens. VM. VM-16893 , Bjarkøy, 68°58′N, 16°42′E, 60–90 m, 07.09.1912, 2 specimens . VM-16894 , Bjarkøy, 68°58′N, 16°42′E, 30–60 m, 06.09.1912, 1 specimen . VM-16895 , Varanger, 70°05′N, 29°30′E, 160–170 m, 18.07.1937, 1 specimen . VM-16896 , Traenfjorden, 66°25′N, 12°15′E, 40 m, 18.07.1939, 1 specimen . VM-16897 , Mefjaera, Vågsfjord, 68°55′N, 16°55′E, 150–200 m, 06.07.1933, 2 specimens . VM- 16898 , Gibostad, 69°21′N, 18°05′E, 40 m, 20.07.1917, 1 specimen (dry) . VM-16899 , Vaerøy, 67°40′N, 12°40′E, littoral, 03.06.1929, 1 specimen . ZMO. B-1287, Håka, Oslofjord, 16.09.1953, 1 specimen . B-1288, Tromsø, 69°40′N, 19°00′E, 2 specimens.

Description

Clathrinidae with a cormus normally composed of a clathroid cormus of regularly and tightly anastomosing tubes and a peduncle. Up to 2 cm in height and 2 cm in diameter. One or several true tubes with a normal choanoderm form the peduncle. Sometimes the peduncle is very short or absent. In specimens without a peduncle the cormus is often narrower at the base than in the apical region. Water-collecting tubes converge at the centre of the sponge, ending in one or sometimes several apical oscula. The single tubes are about 0.3–0.7 mm in diameter. Surface is smooth. Colour white, greyish white or beige when alive, and with almost the same colours in alcohol. Dried specimens usually light beige. The skeleton of the cormus is composed of regular (equiangular and equiradiate) triactines to which parasagittal spicules may be added. The spicules are irregularly orientated in several layers in the walls of the tubes, resulting in a wall thickness of about 80 µm. In the peduncle the skeleton consists of only parasagittal triactines, with the unpaired actine basipetally orientated. All the actines are straight, cylindrical and with blunt tips.

Distribution

The species is found along the entire Norwegian coast from littoral to 500 m depth. G. blanca is the only species among the Norwegian Calcinea that is reported to be common in abyssal depths in the North Atlantic ( Barthel & Tendal, 1993; Janussen et al., 2003). However, recent examination of material from the abyssal Norwegian Sea revealed that these deep records probably represent another new species of Guancha (H. T. Rapp & O. S. Tendal, unpubl. data).

Remarks

In the reticulate/cushion-shaped form the parasagittal spicules are not as numerous, and the unpaired actine is usually shorter than in the pedunculated form. However, parasagittal triactines are always present, also in the young olynthus. The high degree of morphological variation figured by Miklucho-Maclay has previously been questioned ( Burton, 1930a, b). However, material from Trondheimsfjord support some of Miklucho-Maclay’s observations, and the variation is assumed to be mainly due to different life stages of the sponges.

The species has previously been reported from Trondheim and Stavanger ( Burton, 1930b). In his work on the calcareous sponges from the Siboga- Expedition, Burton (1930a) decided to include all species of Guancha (Leucosolenia) with a clathroid cormus (except G. lacunosa ) in Guancha macleayi ( Von Lendenfeld, 1885) , a species originally described from Australia. He retained only Miklucho-Maclay’s olynthus-form as the true Guancha blanca , and the consequence was a cosmopolitan distribution of G. macleayi , a conclusion that has never been supported by later authors. Re-examination of Burton’s specimens from the Norwegian coast revealed that they should be identified as G. blanca .

GUANCHA CAMURA SP. NOV.

( FIGS 17A–C , 18 , TABLE 9)

Type locality: Østerbotn , Porsanger, north-east Norway at 70°15.1′N, 25°21.1′E .

Holotype: TMU-185 , Østerbotn, Porsanger, 70°15.1′N, 25°21.1′E, 27–33 m, 14.08.1956.

Paratype: TMU-184 , Bøkfjord, 69°52.2′N, 29°38.5′E, 50–90 m, 25.06.1937, 1 specimen .

Etymology: From ‘camur’ (Latin), meaning tortuous or bent, referring to the horn-shaped paired actines of the parasagittal triactines from the peduncle.

Additional material examined: 5 specimens.

TMU. TM-186, Jarfjord, 69°40.7′N, 30°24.7′E, 30–45 m, 03.07.1937, 1 specimen . VM. VM-16900 , Evenskjer, 68°35′N, 16°35′E, 60–85 m, 12.08.1913, 2 specimens (dry) . VM-16901 , Gibostad, 69°21′N, 18°05′E, 50 m, 27.07.1917, 1 specimen (dry) . VM-16902 , Lødingen, 68°24′N, 15°58′E, 35–40 m, 18.08.1913, 1 specimen (dry) .

Description

Clathrinidae with a cormus normally composed of a clathroid body of irregularly and loosely anastomosing tubes and a peduncle. Up to 3 cm in height and 2 cm wide. Consistency soft because of the very thin walls of the tubes (about 20 µm). Surface smooth. Several true tubes with a normal choanoderm form the peduncle. The skeleton of the cormus is composed of regular (equiangular and equiradiate) triactines to which parasagittal spicules are frequently added. In the peduncle the skeleton consists of only parasagittal triactines. The unpaired actine of peduncular parasagittal triactines is always basipetally orientated. The paired actines are bent in a manner making them ‘horn’-shaped, whereas the unpaired actine is straight. All actines are cylindrical with slightly blunt tips.

Distribution

The species is found in the northern part of Norway at 27–90 m depth.

The species is also known from Greenland at 125– 200 m depth (my pers. observ.).

Remarks

G. camura sp. nov. differs from other species mainly because of the horn-shaped parasagittal triactines in the peduncle. Such spicules have been found in the olynthus-form of G. ‘ blanca ’ from Neapel ( Metschnikoff, 1879). None of his specimens has been available for examination, but from the distribution of G. camura it is reasonable to believe that they represent two different species. These spicules also bear similarities to sagittal tripods found in G. challengeri ( Polejaeff, 1883) from Australia and New Zealand, but in G. challengeri they are found together with tripods, forming conules on the outer surface. G. challengeri also possesses a solid peduncle ( Polejaeff, 1883), whereas the peduncle is composed of true tubes in G. camura . The species also bears similarities to G. pellucida sp. nov. However, the spicules in G. camura are considerably shorter and thicker than the long and slender spicules in G. pellucida .

GUANCHA LACUNOSA ( JOHNSTON, 1842)

( FIGS 19A–C , 20 , TABLE 10)

Original description: Grantia lacunosa Bean in Johnston, 1842: 176.

Synonyms and citations

Ascandra angulata Von Lendenfeld, 1891 .

Ascandra angulata ( Arnesen, 1901a: 13; 1901b: 68). Ascortis lacunosa ( Haeckel, 1872: 70–71) .

Guancha lacunosa ( Borojevic & Boury-Esnault, 1987: 15–16; Rapp, 1999: 72).

Leucosolenia angulata ( Breitfuss, 1927: 27) . Leucosolenia lacunosa ( Breitfuss, 1933: 241) .

Type locality: Scarborough, UK.

Holotype: According to Burton (1930a) the type is lost.

Previous records: Hjeltefjorden, Bergen ( Arnesen, 1901a).

Material examined: 2 specimens.

VM. VM-16903 , Finnsnes, 69°14′N, 17°58′E, 25–40 m, 18.06.1914 , 1 specimen (dry, peduncle broken off and missing). ZMUB. ZMUB-9678 , Hjeltefjorden, Bergen, 60°35 N, 04°55′E, 55–200 m, 1 specimen .

Description

Sponge composed of an ovoid body of thin, tightly and regularly anastomosing tubes, and a solid peduncle without any choanoderm. Water-collecting tubes converge into one apical osculum. Colour white in ethanol and light beige when dried.

Triactines from the clathroid body range from almost regular to parasagittal with straight actines. Close to the peduncle there are parasagittal spicules with the longest unpaired actine pointing towards the peduncle. The skeleton of the peduncle is composed of large diactines with a break on the middle, and parasagittal triactines with a very long unpaired actine and short paired actines. Irregular diactines of variable sizes are present at low numbers in the peduncle. All actines are cylindrical with slightly blunt to sharp points.

Distribution

From the Norwegian coast, G. lacunosa is only known from Hjeltefjorden in the Bergen area and at Finnsnes in northern Norway, 25–200 m depth.

The species is also reported from the Mediterranean ( Von Lendenfeld, 1891), the Bay of Biscay down to 550 m depth ( Borojevic & Boury-Esnault, 1987), Great Britain ( Ackers et al., 1985) and the Arctic ( Breitfuss, 1911, 1927, 1933; Derjugin, 1915).

Remarks

In his work on the Norwegian sponges from the Norman collection, Burton (1930b) reported Guancha lacunosa from Bergen and Stavanger. Re-examination of these specimens revealed that they should be identified as Guancha blanca and Clathrina nanseni .

GUANCHA PELLUCIDA SP. NOV.

( FIGS 21A–D , 22 , TABLE 11)

Type locality: Porsangerfjord , northern Norway, at approximately 70°15′N, 25°21′E .

Holotype: TMU-179 , Porsangerfjord, approximately 70°15′N, 25°21′E, 140–155 m, 03.08.1931.

Paratype: VM-397 , Senja, 190 m, 08.07.1933, 1 specimen. No more information given on the original label .

Etymology: Named ‘pellucida’ from Latin ‘pellucidus’, meaning translucent, referring to the very thin-walled and translucent tubes of the cormus.

Additional material examined: 4 specimens. IB. Trondheimsfjord, approximately 63°30′N, 10°30′E, 3 different localities, 100–275 m, 1996, 4 small specimens .

Description

Clathrinidae normally composed of a globular and highly compressible clathroid cormus of irregularly but tightly anastomosing tubes, and a very short peduncle. Up to 3 cm in height and 2.5 cm in diameter. Several anastomosing true tubes with a normal choanoderm form the peduncle. In full-grown specimens water-collecting tubes converge into one large apical osculum. Texture soft and fragile. Surface smooth. Colour greyish white and highly translucent when alive, and yellowish white in alcohol. The skeleton of the cormus is composed of 1–3 layers of very slender regular to subregular triactines, to which parasagittal spicules may be added. The walls of the tubes are only 10–15 µm thick. In the peduncle the skeleton consists of parasagittal triactines with unpaired actine of highly variable length. The unpaired actine of peduncular parasagittal triactines is always basipetally orientated. All actines are undulated, especially close to the centre of the spicule, cylindrical or slightly conical with blunt tips.

Distribution: The species is found in the northern part of Norway at 100–275 m depth.

The species is also known from Greenland at 20– 190 m depth and close to Jan Mayen at 890 m depth (my pers. observ.).

Remarks

The species bears some similarities to G. camura sp. nov., but is clearly separated by the long, slender and slightly undulated actines.

Figures

Figure 13. A, fixed specimens of Guancha arnesenae sp. nov. H = holotype. Scale bar = 1 cm. B, parasagittal triactines from the cormus (tc) and from the peduncle (tp). Scale bar = 100 µm.

Figure 14. Distribution of Guancha arnesenae sp. nov. along the Norwegian coast.

Figure 15. A, Guancha blanca at 10 m depth. os = osculum. Scale bar = 1 cm. B, habitus of a fixed specimen. Scale bar = 1 cm. C, section through an ascon tube with mature oocytes (oc). Scale bar = 100 µm. D, slightly parasagittal triactine from the cormus. Scale bar = 100 µm. E, parasagittal triactines from the peduncle. Scale bar = 100 µm.

Figure 16. Distribution of Guancha blanca along the Norwegian coast.

Figure 17. A, holotype of Guancha camura sp. nov. Scale bar = 1 cm. B, triactines from the cormus. Scale bar = 100 µm. C, parasagittal triactines from the peduncle (p). Scale bar = 100 µm.

Figure 18. Distribution of Guancha camura sp. nov. along the Norwegian coast.

Figure 19. A, habitus of a fixed specimen of Guancha lacunosa. Scale bar = 1 mm. B, regular to parasagittal triactines from the cormus (tc) and parasagittal triactines from the peduncle (tp). Scale bar = 100 µm. C, large diactines with an angle in the middle region (ld) and small distorted diactines (sd), all from the peduncle. Scale bar = 100 µm.

Figure 20. Distribution of Guancha lacunosa along the Norwegian coast.

Figure 21. A, holotype of Guancha pellucida sp. nov. Scale bar = 1 cm. B, habitus of a young specimen without a peduncle. os = osculum. Scale bar = 1 mm. C, triactine from the cormus. Scale bar = 100 µm. D, parasagittal triactine from the peduncle. Scale bar = 100 µm.

Figure 22. Distribution of Guancha pellucida sp. nov. along the Norwegian coast.