Leucosolenia qingdaoensis sp. nov. (Porifera, Calcarea, Calcaronea, Leucosolenida, Leucosoleniidae), a new species from China
expand article infoYan-Ling Chu§|, Lin Gong§|, Xin-Zheng Li|§
‡ Institute of Oceanology, Chinese Academy of Sciences, Qingdao, China
§ Center for Ocean Mega-Science, Chinese Academy of Sciences, Qingdao, China
| Laboratory for Marine Biology and Biotechnology, Qingdao, China
¶ University of Chinese Academy of Sciences, Beijing, China
Open Access


A new species of Leucosoleniidae, Leucosolenia qingdaoensis sp. nov., is described. This new species was collected in a scallop-breeding pond from the Yellow Sea and preserved in 75% ethanol. This sponge consists of a dense reticulation of ascon tubes, with the surface minutely hispid and the consistency soft and fragile.

Spiculation of the new species consists of diactines, which are smooth, straight or sometimes slightly curved, triactines of two types, and tetractines with short and curved apical actines; spiculation also slightly overlaps and is somewhat irregularly assembled. Together these form a thin layer of skeleton, with a small number of cells, which results in a transparent, white sponge. As a typical asconoid feature, all internal cavities of the sponge are lined with choanocytes, and there is no fully developed inhalant system. Comparisons with other Leucosolenia reported from the Pacific Ocean are also made.


Sponge, taxonomy, Yellow Sea


The family Leucosoleniidae is characterised by a branched and rarely anastomosed cormus and asconoid aquiferous system; there is neither a common cortex nor a delimited inhalant or exhalant aquiferous system (Minchin 1900). The family includes three genera (Borojevic et al. 2002): Ascyssa Haeckel, 1872, Ascute Dendy & Row, 1913, and Leucosolenia Bowerbank, 1864. They can be easily distinguished by their skeletons: the skeleton of Ascyssa contains only diactines; the skeleton of Ascute exhibits giant longitudinal diactines forming a continuous layer on the external surface, and includes triactines and tetractines; and the skeleton of Leucosolenia lacks any of these obvious characteristics in the above two genera. Instead, the skeleton of Leucosolenia is characterised by being composed of diactines, triactines and/or tetractines, without a reinforced external layer on the tubes.

The genus Leucosolenia comprises 40 living species worldwide (Van Soest et al. 2019), of which only three species, L. microspinata Longo, 2009, L. salpinx Van Soest, 2017, and L. parthenopea Sarà, 1953, were named after 1950; 11 species were described by Haeckel between 1870 and 1872. The literature of this genus is relatively old, and the descriptions contained therein of the species of Leucosolenia were simple, almost without details and illustrations of the body shapes and spicules. Thus, a taxonomic revision of this genus is very difficult, and to date, no worldwide revision of the genus has been made.

The localities of the 15 known species of Leucosolenia recorded from the Pacific Ocean are shown in Figure 1. Seven species (L. eleanor Urban, 1906, L. minuta Tanita, 1943, L. mollis Tanita, 1941, L. pyriformis Tanita, 1943, L. serica Tanita, 1942, L. tenera Tanita, 1940, and L. ventosa Hôzawa, 1940) were reported from the Japanese waters (Sagimi Sea, Wakayama Prefecture, Onagawa Bay, Mie Prefecture, Matsushima Bay, Izushima, Wagu Miye Prefecture, respectively). Leucosolenia macquariensis Dendy, 1918 was reported from the west coast of Macquarie Island; L. australis Brøndsted, 1931 was reported from Comau Fjord; L. albatrossi Hôzawa, 1918 was reported from Copper Island and the Komandorski Islands; L. echinata Kirk, 1893 and L. rosea Kirk, 1896 were reported from New Zealand; L. lucasi Dendy, 1891 was reported from Port Phillip Heads, Australia; L. nautilia Laubenfels, 1930 was reported from California, USA; and L. feuerlandica Tanita, 1942 was reported from Tierra del Fuego, South America. The Leucosolenia species reported from the coasts of Japan account for most species. The type specimens of new species were found in the Yellow Sea, very close to Japan.

Figure 1. 

Distribution of Leucosolenia A location in the Pacific Ocean B detail of the type locality in the Japanese coast: (1) Komandorski Islands (L. albatrossi Hôzawa, 1918); (2) Comau Fjord (L. australis Brøndsted, 1931); (3) Cook Strait, Poverty Bay, Kawakawa (L. echinata Kirk, 1893); (4) Francisco Bay, California; Sukumo ôsima, Kôti Prefecture, Sagimi Sea (L. eleanor Urban, 1906); (5) Tierra del Fuego (L. feuerlandica Tanita, 1942); (6) Port Phillip Heads, Australia, and New Zealand (L. lucasi Dendy, 1891); (7) Macquarie Island (L. macquariensis Dendy, 1918); (8) Wakayama Prefecture (L. minuta Tanita, 1943); (9) Onagawa Bay (L. mollis Tanita, 1941); (10) Monterey Bay, California (L. nautilia Laubenfels, 1930); (11) Mie Prefecture (L. pyriformis Tanita, 1943); (12) New Zealand (L. rosea Kirk, 1896); (13) Yodomi, Sagami Sea (L. serica Tanita, 1942); (14) Matsushima Bay, Onagawa Bay, Izushima (L. tenera Tanita, 1940); (15) Wagu Miye Prefecture (L. ventosa Hôzawa, 1940); (*) Qingdao (L. qingdaoensis sp. nov.).

Materials and methods

The specimens were collected in a scallop-breeding pond from the Yellow Sea and were preserved in 75% ethanol. Two specimens were deposited in the Marine Biological Museum of the Institute of Oceanology in the Chinese Academy of Sciences (IOCAS), Qingdao, China.

For examination of the spicules, a small piece of specimen was cut and placed in a 1.5 mL microcentrifuge tube to which 1000 µL of sodium hypochlorite solution was added (Kersken et al. 2016). The mixture was then vortexed, placed at environmental temperature, and vortexed occasionally during incubation until it was completely lysed. Next, the sample was centrifuged at 8000 rpm for 2 min, the supernatant was poured off, 1000 µL of distilled water was added, and the sample was again centrifuged at 8000 rpm for 2 min. This procedure was repeated four times, then the spicules were washed three times with 96% ethanol and then the spicules were preserved in the third ethanol solution.

Scanning Electron Microscopy (SEM) was performed with a Hitachi S3400N. Preserved spicules for SEM were adhered to stubs with double-sided carbon conductive tape and coverslip. After dehydration, the spicules were coated with gold in a Hitachi MC1000 (LOPES 2018).

Measurements of at least 20 spicules of each type were performed using an optical microscope (Nikon Eclipse Ni) with a micrometric eyepiece. The length from the tip to the base and the thickness at the base of each actine were measured. The reported numbers refer to the range of measurements for each spicule type. Photographs were taken with a stereomicroscope (Zeiss Stemi 2000-c) and an optical microscope (Nikon Eclipse Ni-U) equipped with a digital camera to evaluate difference between the length of the unpaired and paired actines of each type of triactine. For comparison with the new species, we only selected those species of Leucosolenia reported from the Pacific Ocean.



Class Calcarea Bowerbank, 1862

Subclass Calcaronea Bidder, 1898

Order Leucosolenida Hartman, 1958

Family Leucosoleniidae Minchin, 1900

Genus Leucosolenia Bowerbank, 1864

Leucosolenia qingdaoensis sp. nov.

Figs 1, 2, 3, 4; Tables 1, 2

Type material

Holotype : MBM181606, scallop-breeding pond on southeastern Shandong Peninsula, China, June 1988, 0–0.3 m depth, collected by Shue Li, 35°58'N, 120°11'E. Paratype: MBM181476, Zhonggang, Qingdao, China, 7 June 1984, 0–0.6 m depth, 36°06'N, 120°21'E.

Type locality

Qingdao, Yellow Sea.


The name is derived from the type locality, Qingdao, China.


The sponge is arborescent, consisting of many thin-walled tubes, which are copiously ramified but never anastomosed. The sponge occurs as growth form. The oscula are terminal on erect tubes. The color of the sponge is white after being preserved in alcohol and in vivo. The external walls of the tubes are hairy, with diactines protruding at right or oblique angles from the body; the surface is minutely hispid, and the consistency is soft and fragile. The holotype measures 21.32 × 3.38 mm (height × width). The wall of the sponge body is very thin, and there is no fully developed inhalant system, the gap between the skeleton and the cell on the wall arrange evenly (Fig. 2F); only a small amount of cells is distributed on the thin sponge skeleton (Fig. 2C–F), which is a typical asconoid feature. All internal cavities of the sponge are lined by choanocytes.

Figure 2. 

Leucosolenia qingdaoensis sp. nov. A holotype B paratype C detail of oscula (stereo microscope) D detail of root-like structures (stereo microscope) E detail of oscula (optical microscope) F detail of root-like structures (optical microscope); arrowhead pointing at the ostium.

Skeletal arrangement

The skeleton consists of multifarious diactines, sagittal triactines of two types, sagittal tetractines with bent apical actines and triactine-like basal actines; together these form the wall of the ascon-type sponge body.

In the apical osculum (Fig. 2C, E), there are paired actines of triactines and tetractines, some additional tangential diactines, together forming a clear line dividing the apical oscula, and some radial diactines projecting beyond the apical osculum with different length.

In the sponge body (Fig. 2C, E), the triactines and tetractines are regularly arranged, their paired actines are parallel to the apical oscula, and the unpaired actines point downward, with slight folding allowed, but never overlapping; in contrast to the triactines and tetractines, the diactines are arranged more irregularly but generally point downward.

In the root-like structures (Fig. 2D, F), the arrangement of triactines and tetractines is the same as that in the body, but the arrangement of diactines is different; most of them tangentially project beyond the surface, which results in the surface having a slightly hispid appearance.

By observing the sponge tissue taken from different parts, it is clear that as the diameter of the tubes decreases, the contents of small diactines and small triactines increase. This observation can suggest that in the growth zone spiculogenesis is more intense.


Diactines. There is only one type of diactine (Fig. 3A1–3), though the diactines vary in size and shape, their width varies from 24 µm to 61 µm, the length of diactines vary from 43 µm to 421 µm but half of the diactines present a length of 200–300 µm (Fig. 4). The shapes of the diactines are straight or slightly curved in different directions. The variation in Leucosolenia is very common and considerable.

Triactines. Two types of triactines are present, with actines straight or undulated. Their ends are generally sharp or asymmetrical (Fig. 3B12). The paired actines are slightly curved. Some deformations are present.

Type 1: triactines with paired actines longer than unpaired actines (Fig. 3B1): unpaired actines 42–105 × 3–5 µm; paired actines 63–105 × 3–5 µm.

Type 2: triactines with unpaired actines longer than paired ones (Fig. 3B2): unpaired actines 76–129 × 3–4 µm; paired actines 60–104 × 3–4 µm.

Tetractines. A relatively small number of tetractines are observed, approximately 10 per 100 spicules, with straight and fusiform actines (Fig. 3C12). The tetractines are similar to triactines but with the addition of apical actines, the apical actines are fairly stout and short, sharply pointed and curved: unpaired actines 93–119 × 2–5 µm; paired actines 50–93 × 2–5 µm; apical actines 11–29 × 2–5 µm.

Figure 3. 

Spicules of Leucosolenia qingdaoensis sp. nov. (holotype) A1–3 = diactines; B1 = triactines of type 1; B2 = triactines of type 2; C1–2 = tetractines.

Figure 4. 

Leucosolenia qingdaoensis sp. nov. Size-class distribution of diactines (holotype).


Three species described by Tanita (L. minuta, L. pyriformis, and L. serica) exhibit only regular (equiangular and equiradiate) spicules. This characteristic does not fit the description of Leucosolenia, L. qingdaoensis sp. nov. can be easily differentiated from the 12 species of Leucosolenia reported from the Pacific Ocean. The skeletal compositions of these species are shown in Table 1.

Spicules dimensions of Leucosolenia Bowerbank, 1864 in the Pacific Ocean. Measurements are reported in µm.

Triactines Tetractines Diactines References
Unpaired Paired Unpaired Paired Apical
Length/Width Length/Width Length/Width Length/Width Length/Width Length/Width
L. albatrossi 70–90/8 80–100/8 70–90/8 80–100/8 40–60/6 70–90/8 Hôzawa 1918
60–90/8 130–240/8 60–90/8 130–240/8 40–60/6–8
L. australis 69–122/6 66–106/6 66–119/6 69–99/7 27–41/4 41–49/1 Azevedo et al. 2009
L. echinata 100/10 130/10 130/15 150/15 70/15 240–730/10–5 Kirk 1893
L. eleanor 80/7 80/7 140/9 140/9 140/9 105/4 Laubenfels 1932
140/7 140/7 434/9
L. feuerlandica 50–70/12–18 70–95/12–18 60–70/8–10 75–90/8–10 40–50/6–8 70–90/4–6 Tanita 1942
60–70/8–10 75–90/8–10
L. lucasi 100/5 70/5 100/5 70/5 <70/5 160/5 Dendy 1891
L. macquariensis 980/9 980/9 980/9 980/9 140/6 Dendy 1918
L. minuta 130–175/14–18 130–175/14–18 60–75/8–10 60–75/8–10 50–60/7–10 Tanita 1943
60–75/8–10 60–75/8–10
L. mollis 70–130/6–8 90–140/6–8 70–130/6–8 90–140/6–8 35–55/6 230–400/7–10 Tanita 1941
L. nautilia 140/9 140/9 140/9 140/9 30/8 400/10 Laubenfels 1932
L. pyriformis 180–190/12–18 180–190/12–18 180–190/12–18 180–190/12–18 150–260/8–15 630–800/40–55 Tanita 1943
L. rosea 300/70 300/70 140/10 140/10 110/8 Kirk 1896
200/18 200/18
L. serica 140–210/7–8 140–210/7–8 140–210/7–8 140–210/7–8 90–135/8–10 Tanita 1942
L. tenera 80–180/7–10 90–210/7–10 80–180/7–10 90–210/7–10 30–10/6–8 200–530/8–12 Tanita 1940
L. ventosa 100–120/10 85–100/10 Hôzawa 1940
150–180/20–25 140–150/20–25
100–120/10–14 70–90/10–14
L. qingdaoensis sp. nov. 42–104/3–5 63–105/3–5 93–119/2–5 50–93/2–5 11–29/2–5 43–422/4–7 Present paper
76–129/3–4 60–104/3–4

The new species exhibits one type of diactine. In L. ventosa and L. rosea, there is no record of diactines, and in L. mollis and L. nautilia, there are two types of diactines. The triactines of L. ventosa are 2–8 times thicker than those in the new species; the triactines of L. rosea are 10–35 times thicker than in the new species; and L. mollis only has one type of triactine and all rays being nearly equally thick. The diactines of L. nautilia are extremely large, with a length of 1 mm and a thickness of 20 µm (Laubenfels 1932), while in the new species the diactines are less than 8 µm thick. Laubenfels (1932) gave few details on the actines, but L. nautilia differs from the new species by having only one type of triactine.

The difference between L. albatrossi and the new species is obvious. The diactines of L. albatrossi are club-shaped, while the diactines of the new species are spindle-shaped.

The sagittal triactines of the new species distinguish it from L. macquariensis, L. tenera, and L. eleanor. The new species have two types of sagittal triactines, while L. macquariensis and L. tenera only have one type of sagittal triactine, with rays of approximately equal length. Leucosolenia eleanor have both sagittal and regular triactines.

The new species, with slender and long diactines, the longest diactines 5 times longer than those of L. feuerlandica, is distinct from that species. Additionally, the triactines of the new species are sagittal, and the actines straight or undulated. However, the triactines of L. feuerlandica are pseudoderm sagittal and are tripod-shaped.

Leucosolenia echinata, L. lucasi, and L. qingdaoensis sp. nov. have many features in common, including their body shape, colour in alcohol, general arrangement, shape of diactines, and apical ray, but they show important differences in the shape of their triactines. The new species has two types of triactines; L. lucasi and L. echinata only have one type of triactine. The triactines of L. lucasi are sagittal, but the three angles are roughly equal; the triactines of L. echinata are generally regular, and frequently slightly sagittal, with the oral angle largest and the basal ray longest.

Spicules measurements of Leucosolenia qingdaoensis sp. nov. (holotype).

length(µm) width(µm)
min mean max sd min mean max sd n
Diactines 43 219 422 93 1 4 7 1.7 50
Triactines 1
paired 63 83 105 9 3 4 5 0.8 50
unpaired 42 66 105 13
Triactines 2
paired 60 79 104 11 3 3 4 0.4 50
unpaired 76 102 129 15
paired 50 77 93 12 2 4 5 0.8 20
unpaired 93 104 119 11
apical 11 21 29 6

Key to the species of Leucosolenia in the Pacific Ocean

1 Skeleton contains only regular spicules 2
1a Skeleton contains sagittal spicules 4
2 Skeleton including diactines L. pyriformis
2a Skeleton without diactines 3
3 Rays are stout L. minuta
3a Rays are relatively thin L. serica
4 Skeleton contains diactines, triactines, and tetractines 5
4a Skeleton contains triactines and tetractines L. rosea
4b Skeleton contains only triactines L. ventosa
5 Skeleton contains one type of diactine 8
5a Skeleton contains two types of diactines 6
6 Diactines are club-shaped L. macquariensis
6a Diactines are spindle-shaped 7
7 Skeleton without large diactines L. mollis
7a Skeleton including large diactines L. nautilia
8 One tip of diactines has spines L. australis
8a Diactines have no spines 9
9 Skeleton contains one type of triactine 10
9a Skeleton contains two types of triactines 11
10 Sagittal triactines with rays are of approximately equal in length L. tenera
10a Sagittal triactines with rays are of different lengths L. lucasi
10b Triactines are generally regular, slightly sagittal L. echinata
11 Skeleton including tripod type of triactines L. feuerlandica
11a Skeleton without tripod type of triactines 12
12 Diactines have one ‘lance head’ type ends L. albatrossi
12a Diactines have two smooth and sharply pointed ends 13
13 Skeleton contains both sagittal and regular triactines L. eleanor
13a Skeleton contains only sagittal triactines L. qingdaoensis sp.nov.


This work was supported by the National Natural Science Foundation of China (no. 41706188), the Science and Technology Basic Work Program (2014FY110500), Biological Resources Programme, Chinese Academy of Sciences KFJ-BRP-017-37, and the Scientific and Technological Innovation Project Financially Supported by Qingdao National Laboratory for Marine Science and Technology (no. 2015ASKJ01). We thank Dr Yuanyuan Sun for assistance with SEM operations, and we appreciate Dr Qinghe Liu for the help with photography. We also thank Mr Hongfa Wang, Dr Lin Ma, Dr Jixing Sui, and Mr Youwei Tzeng for providing important references needed for this study.


  • Borojevic R, Boury-Esnault N, Manuel M, Vacelet J (2002) Order Leucosolenida Hartman, 1958. In: Hooper JNA, Van Soest RWM (Eds) Systema Porifera: A Guide to the Classification of Sponges. Kluwer Academic/Plenum Publishers, New York, 1157–1184.
  • Brøndsted HV (1931) Die Kalkschwämme der Deutschen Südpolar Expedition 1901–1903. Deutsche Südpolar-Expedition, 1901–03 20: 1–47.
  • Dendy A (1891) A monograph of the Victorian sponges, I. The organisation and classification of the Calcarea Homocoela, with descriptions of the Victorian Species. Transactions of the Royal Society of Victoria 3: 1–81.
  • Dendy A (1918) Calcareous sponges. Scientific Reports of the Australasian Antarctic Expedition, 1911–1914, Series C, Zoology and Botany 6(1): 1–17.
  • Hôzawa S (1918) Reports on the calcareous sponges collected during 1906 by the United States Fisheries Steamer Albatross in the Northwestern Pacific. Proceedings of the United States National Museum 54: 525–556.
  • Hôzawa S (1940) On some calcareous sponges from Japan. Science Reports of the Tohoku Imperial University 15: 29–58.
  • Kersken D, Feldmeyer B, Janussen D (2016) Sponge communities of the Antarctic Peninsula: influence of environmental variables on species composition and richness. Polar Biology 39(5): 851–862.
  • Laubenfels MWde (1930) The sponges of California (abstracts of dissertations for the degree of doctor of philosophy). Stanford University Bulletin 5(98): 24–29.
  • Lopes MV, Padua A, Condor-Lujan B, Klautau M (2018) Calcareous sponges (Porifera, Calcarea) from Florida: new species, new records and biogeographical affinities. Zootaxa 4526(2): 127–150.
  • Minchin EA (1900) Sponges. A treatise on zoology 2: 1–178.
  • Tanita S (1940) Calcareous sponges of Matsushima Bay. Science Reports of the Tôhoku Imperial University 4(15): 165–177.
  • Tanita S (1941) Calcareous sponges obtained from Onagawa Bay and its vicinity. Science Reports of the Tôhoku Imperial University 16(3): 263–282.
  • Tanita S (1942a) Report on the calcareous sponges obtained by the Zoological Institute and Museum of Hamburg. Part II. Science Reports of the Tôhoku University 4(17): 105–135.
  • Tanita S (1942b) Calcareous sponges collected in the Kanto District, Japan. Science Reports of the Tôhoku Imperial University 17(1): 17–69.
  • Tanita S (1943) Studies on the Calcarea of Japan. Science Reports of the Tôhoku Imperial University 17: 353–490.
  • Urban F (1906) Kalifornische Kalkschwämme. Archiv für Naturgeschichte 72 (I): 33–76. [pls VI–IX: 36–55]
  • Van Soest RWM, Boury-Esnault N, Hooper JNA, Rützler K, de Voogd NJ, Alvarez B, Hajdu E, Pisera AB, Manconi R, Schönberg C, Klautau M, Picton B, Kelly M, Vacelet J, Dohrmann M, Díaz M-C, Cárdenas P, Carballo JL, Ríos P, Downey R (2019) World Porifera Database. Leucosolenia Bowerbank, 1864. World Register of Marine Species. [Accessed on: 2019-05-07]