Corresponding author: Rob Van Soest(
Academic editor: Roberto Pronzato
We present a review of astrophorid species possessing calthrops megascleres as structural megascleres (including species with dichotriaene modifications, but excluding mesotriaene and trichotriaene bearing species). Radiating oxeas characteristic of most astrophorids are lacking in such sponges, but auxiliary oxeas are apparently present in some species. These sponges are currently assigned to two families,
The present study addresses the taxonomy of some genera of the order
A further purpose of this paper is to propose a slightly altered generic allocation of species, facilitating easier recognition of the morphological classification, but the familial allocations will be left as they are in the Systema Porifera (
The
The monospecific genus
The calthropellid
We will not include in our study the pachastrellid genus
Several species of
Species with structural oxeas in a radial position are here excluded and referred to other astrophorid genera.
A recent paper by
Below, we review the species of the (sub-)genera
Specimens in the collections of the National Centre for Biodiversity (formerly Zoological Museum of the University of Amsterdam (
Specimens were studied from thick sections cut at right angles to the surface and from dissociated spicules, using light microscopy and a JEOL Scanning Electron Microscope. Digital images of the spicules were assembled on a black background and aligned and cleaned up using Adobe Photoshop CS3.
The left over part of the spicule suspension was used for light microscopy measurements. Measurements of megascleres include for calthrops cladus length × width, cladome size, i.e. the distance between the apex of a cladus and the imaginary line between two opposing cladi endings; for dichocalthrops, protocladus length × width, deuterocladus length × width, rhabdus length × width and cladome size. Measurements of microscleres include for sanidasters length × width including the spines, for toxa-like microscleres length × width, for euasters greatest diameter including rays. Unless otherwise stated, minimum-
In the descriptions and definitions, the word calthrops will be used in singular and plural sense (in accordance with
1 | Megascleres include trichotriaenes | |
– | No trichotriaenes | 2 |
2 | Megascleres include meso-dichotriaenes; microscleres include streptasters | |
– | Megascleres do not include meso-dichotriaenes | 3 |
3 | Microscleres include streptasters (amphiasters, spirasters, metasters) | |
– | Microscleres sanidasters, ataxasters or euasters; no streptasters | 4 |
4 | Microscleres include sanidasters, no euasters | |
– | Microscleres include euasters in some form; no sanidasters |
Phylum
Class
Order
Family
Many authors have pointed out the similarities of
Calthrops are characteristically variable in size (cladi measuring from 25 – 648 µm), and shape, with conical-straight, curved, stunted, deformed, bifid cladi, frequently one of the cladi being longer than the others, sometimes lacking one (‘tripods’) cladus or with one or more reduced cladi, occasionally with five or more cladi. Short-shafted dichotriaenes (dichocalthrops) are present in twelve of the currently named and recognized species, and in five of these have replaced the calthrops entirely. When both megasclere types are combined in a single species, dichocalthrops are often distinctly smaller than the calthrops and the proportion of the two varies among individuals of the species.
Natural products: three unrelated compounds with biological actitivity, such as anti-tumor or antibiotic activity, have been extracted from species of
.
1 | Toxa-like microscleres present |
|
– | No toxa-like microscleres | 2 |
2 | Sanidasters divisble in two categories, thin-and-long and thick-and-short, the latter often compressed to form pseudasters |
|
– | Sanidasters may be variable but not clearly divisible in shape categories, no pseudasters |
|
Key to the subgenera of the genus
.
1 | Robust toxas, up to 100 µm in length |
|
– | Thin toxas only up to 70 µm in length |
Schizoholotype (2 slides, one with dissociated spicules, the other with section at right angles to surface): NHM collection, Bwbk. 542 (labeled as
Holotype of
(amalgamated from various descriptions of material from the British Isles and NW France). Cushion-shaped to massive-lobose (
Skeleton: Main skeleton of confused, randomly arranged calthrops, a layer of sanidasters and many toxas occur near the surface, but these also occur in the choanosome. There are many large pigment cells (?) with black inclusions evident in sections.
Spicules: Calthrops, toxiform microscleres, sanidasters.
Calthrops (
Smooth toxiform microscleres (
Sanidasters (
Spicule data reported for specimens of
Author | calthrops cladi | cladome | sanidasters | toxas | locality |
---|---|---|---|---|---|
81–318 × 24–41 | 120–480 | 22–31 × 0.5–7 | 64–111 × 1–2 | Devon Guernsey | |
108–356 × 17–37 | 142–472 | 21–31 | 51–103 | Devon, UK | |
|
310–320 × 38–45 | 465–480* | 21–27 × 6–8 | 75–90 × 2–3 | Bretagne |
|
95–210 × 30 | 142–310* | not reported | not reported | Asturias |
|
58–119 ×5–16 | 126–186 | 17–22 × 0.5–3 | 37–58 × 0–1 | Gorringe |
Berlengas unpubl. | 75–165 × 7–27 | 110–240 | 16–24 × 1.5–2 | 42–69 × 0.5–1.5 | Lagosteira |
Portuguese main coast unpublished | 118–135 × 12–22 | 175–200 | 19–21 × 2 | 48–51 × 0.5–1.5 | Arabida |
In crevices in vertical rock faces in clean water. Particularly common on limestone substrata. From the extreme low water mark to a few meters subtidally in rock pools and caves.
British Isles (SW coast of England and N and W coast of Ireland), and France (Roscoff, Iles de Glénan). There is a record from the northern Gulf of Mexico (
Named after Mrs. Buckland of Guernsey, who collected one of the type specimens (in fact the species should probably have been named “bucklandae” since the named person is female).
Bowerbank’s type specimen (a fragment, BMNH 1877.5.21.142,
Carter’s type specimens (
In view of some of the discrepancies we considered it worthwhile to investigate whether spicular differences of specimens within the known range of the species would yield a pattern that could explain some of the differences. We compared spicular data from the northern samples cited above with those of samples from areas in the southern part of the range, from the coasts and offshore localities of Portugal obtained from Dr Joana Xavier and examined spicule sizes and shapes. This led us to the conclusion that there are consistent differences between samples from these areas which should be recognized at the subspecies level.
Holotype ZMA Por. 21810, Portugal, Gorringe Bank, Gettysburg Peak, 31–38 m, coll. J. Xavier, 2006 (cf.
Additional specimens. Xavier collection, field nr. B05.09.36, Portugal, Berlengas Archipelago, Lagosteira, 7 m, 16–IX–2005, coll. J.R.B.T. Xavier.
Xavier collection, field nr. B05.09.59, Portugal, Berlengas Archipelago, Lagosteira, 6–7 m, 16–IX–2005, coll. J. Cristobo.
Xavier collection, Portugal, field nr. B05.09.98, Berlengas Archipelago, Lagosteira, 6 m, 18–IX–2005, coll. J.R.B.T. Xavier.
Xavier collection, field nr. B05.09.267, Portugal, Berlengas Archipelago, Gruta do Carreiro Maldito, 6–8 m, IX–2005, coll. A. Cunha.
Xavier collection, Portugal, field nr. A03/73, Arabida, Ponta da Passagem, 7 m, 16–VII–2003, coll. J.R.B.T. Xavier.
(
Skeleton: a layer of microscleres overlying a loose mass of calthrops megascleres. Spicular density appears lower than in the specimens from the British Isles.
Spicules: calthrops, toxas, sanidasters.
Calthrops (
Toxa-like microscleres (
Sanidasters (
Encrusting, typically bridging crevices and gaps in the substrate, sublittoral down to 6–38 m.
Portuguese main coast, Berlengas Islands, Gorringe Seamount. Furthermore, there are reports from the north coast of Spain (Ferrer-Hernandez, 1918; Babio & Gondar, 1978 (not seen);
The spicule measurements of the present specimens are generally significantly smaller than those provided by most previous authors (
The toxa–like microscleres of
.
1 | Megascleres exclusively calthrops, no proper dichocalthrops (but bifid cladi may be present in the calthrops) | 2 |
– | Megascleres exclusively dichocalthrops, no proper calthrops | 8 |
– | Megascleres including both dichocalthrops and calthrops | 10 |
2 | Longest cladi of calthrops less than 200 µm | 3 |
– | Longest cladi over 200 µm | 5 |
3 | Longest cladi of calthrops less than 100 µm | 4 |
– | Longest cladi of calthrops up to 200 µm | |
4 | Calthrops exclusively three-claded, may be absent; sanidasters up to 20 µm |
|
– | Calthrops both normal and three-claded, always present; sanidasters 8–12 µm |
|
5 | Longest cladi of calthrops over 250 µm (maybe up to 400 µm) | |
– | Longest cladi of calthrops less than 250 µm | 6 |
6 | Sanidasters up to 25 µm long | |
– | Sanidasters only 10–15 µm long | 7 |
7 | Live colour red (dark in preservation) |
|
– | Live colour yellow (white in preservation) | |
8 | Cladome only up to 230 µm in diameter |
|
– | Cladome up to 450 or more µm in diameter | 9 |
9 | Insinuating in calcareous algae, colour brownish |
|
– | Encrusting, white | |
10 | Calthrops very large, cladi up to 648 µm |
|
– | Cladi of calthrops less than 250 µm | 11 |
11 | Calthrops with one cladus distinctly longer | |
– | Cladi may be unequal in length but none are distinctly longer | 12 |
12 | Dichocalthrops distinctly smaller than calthrops | 13 |
– | Dichocalthrops similar in cladome size as calthrops | 14 |
13 | Sanidasters with low short spines |
|
– | Sanidasters profusedly spined | |
14 | Largest dichocalthrops cladomes over 300 µm | |
– | Largest dichocalthrops cladomes less than 300 µm | 15 |
15 | Sanidasters fusiform, pointed | 16 |
– | Sanidasters rhabd-like, blunt | 17 |
16 | Colour pink, thickness of cladi of megascleres 10 µm |
|
– | Colour grey, thickness of cladi of megascleres up 27 µm |
|
17 | Oxeas present |
|
– | No oxeas |
BMNH 1931.1.1.31a, Invisible Bank, Andaman Islands, slide only of which it is uncertain whether it represents the present species. Type: Indian Museum Calcutta ? (not seen).
(from Carter and Sollas). “Sponge excavating. Spicules: Megasclere dichotriaene, rhabdome 0.210 × 0.042 mm. Microsclere rod-like with numerous small spines 0.0127 mm in length. Habitat: Gulf of Manaar”.
The exact properties and variation of this species have not been established so far, so we can only provide a discussion pending a proper revision. The type material has not been reexamined and its precise whereabouts are uncertain.
In contrast to previous descriptions,
If all these specimens are members of a single species, then it occurs on both sides of the Gulf of Bengal as well as on the islands in the middle of it and to the east into Indonesia. However, specimens need to be reexamined.
Holotype MNHN DT 3635, labeled “
Additional specimen ZMA Por. 15101a, Banyuls, ‘coralligène’, 42.4833°N; 3.138°E, 10 m, coll. S. Groot, 10 August 1981.
The wet holotype (
Spiculation of the holotype (
Spicule data reported for specimens of
a. Specimens from the Northeastern Atlantic-Mediterranean area | ||||
---|---|---|---|---|
Author | calthrops | dichocalthrops | sanidaster | locality |
41–188 × 3–29 cladome 57–252 | prot. 20–28 deut. 15–36 rhabd 28–57 cladome 67–105 | 11–19 × 1–2 | Algeria | |
36–183 × 4–28 cladome 62–258 | prot. 14–21 deut. 21–33 rhabd 36–45 cladome 73–93 | 11–18 × 1–2.5 | Algeria | |
|
60 cladome 90* | present (rare) | 8.3 | Algeria |
|
60–80 × 8 cladome 90–120* | present (small) | 6–7 × 1.5 | Adriatic |
|
170–200x 25–30 cladome 255–300* | present | 12–15 × 2–3 | Banyuls |
|
50–238x 18–35 cladome 75–357* | prot.15–18 deut. 22–100 rhabd 50–120 cladm 72–218* | 10–16 × 2–3 | Adriatic |
|
60–250 × 30 cladome 90–375* | prot.25–30 deut. 20–70 rhabd 40–80 cladome 80–190* | 12–13 | Portugal (Gettysburg) |
|
40–60 (rare) cladome 60–90* | prot.not given deut.not given rhabd not given cladm 60–210 | 10–20 | Italy |
|
40–200 cladome 60–300* | prot.20–40 deut.25–50 rhabd not given cladm 80–180* | 12–16×2.5 | Italy |
|
30–206×3–23 cladome 45–309* | cladi 44–65 rhabd 75–80 cladm 80–125* | 11–18 × 0.7–1.5 | Azores |
ZMA 15101 unpublished | 56–195 × 7–20 cladome 92–238 | proto 24–31 deut. 15–21 rhabd. 35–40 cladome 78–98 | 16–22 × 1.5–2 | Banyuls |
b. Specimens described as different species suspected to be D. plicatus. | ||||
92–165 × 14–31 cladome 180–210 | proto 16–18 deut. 15–32 rhabd 45–50 cladome 60–74 | 11–18 × 2–3 | Adriatic | |
45–175 × 5–21 cladome 67–263* | cladome 77–102 | 8–15 × 0.7–4 | Naples | |
c. Specimens originally identfied as D. plicatus here assigned to new species. | ||||
92–426 × 8–55 cladome 126–618 | not present | 11–17 × 1–4 | Senegal | |
36–480 × 6–60 cladome 61–810 | not present | 12–21 × 1.5–3 | Mauritania | |
|
250 × 30 µm | not present | 10–15 × 2 | Senegal |
ZMA 21697 so far unpublished | 141–295 ×11–48 cladome 228–475 | not present | 13–21 × 1–2 | Aegean Sea |
not present | prot. 42–56 deut. 23–204 rhabd. 71–252 cladome 132–475 | 11–16 × 1–2 | Cape Verde Is. | |
96–258 × 198–37 clad.186–420 | prot. 19–30 deut. 55–192 rhabd. 58–132 clad. 129–361 | 15–21 × 3–4 | Fiji | |
d. Specimens from the Indo-West Pacific suspected to belong to undescribed species | ||||
100–240 × 15–27 cladome 150–360* | not present | 12–25 × 2–3 | Malaysia | |
MSNG MD6,MD2 |
70–90 cladome rare | prot.12–25 105–135*deut. 20–82 rhabd. 41–102 cladm.60–200* | 8–13 × 2–3 | Maldives |
Calthrops (
Dichocalthrops (
Sanidasters (
Additionally, we observed some oxeas of uniform size, approximately 150 × 3–4 µm, assumed to be foreign to the sponge.
Description of the BMNH Schmidt’s type slide (
The only non-type specimen available to us, ZMA Por. 15101 from Banyuls has spiculation closely similar to the type material (
Encrusting and insinuating in crevices, large depth range down to 100 m.
Originally reported from Algeria. Elsewhere reported with certainty from Banyuls, Naples and the Adriatic. Possibly some of the records from the adjacent North Atlantic (
Spicule dimensions of reported specimens are presented in
We believe alleged records of this species outside the Mediterranean (
Several species of
Syntype fragment ZMB 2409, dredged near Lesina, Croatia.
Encrusting and insinuating among stones and coralline algae, with oscular elevations. The fragment examined by us was a thin rounded papilla-like extension (probably from the larger of the two type specimens described by Lendenfeld). Consistency slightly rubbery. Colour (alcohol) orange.
Skeleton: at the surface consisting of a dense crust of microscleres, subdermally the skeleton consists of a mass of calthrops, embedded in a fibrous-organic groundmass.
Spicules: calthrops, dichocalthrops and sanidasters, no complete oxeas are found in the fragment examined, but a few broken monaxones were present.
Calthrops (
Dichocalthrops (
Sanidasters (
No data.
Adriatic. So far there is only a single record of this species.
A major difference with the original description is the lack of giant oxeas (4000 × 70 µm), some centrotylote, reported and pictured by
This species is similar in most respects to
None (no response from an online form request to the Stazione Zoologica, Naples).
(from
Skeleton: no data.
Spicules: calthrops, dichocalthrops, sanidasters, oxeas (but see discussion).
Calthrops with the fourth cladus often shorter, occasionally triactines, cladi 45–175 × 5–21 µm.
Dichocalthrops, cladome 77–102 µm, thickness of cladi 3.5–6 µm, no individual proto-, deuteroclad or rhabd measurements provided.
Sanidasters, amphiaster-like with spines concentrated on both sides, length 8–15 µm, thickness 3–4 µm (with spines) 0.7–1.5 µm (without spines).
Oxeas, extreme size variation, 65–930 × 1.8–21 µm, no indication what their structural position is within the sponge.
Holotype ZMA Por. 21530, off the coast of Senegal, coll. F.P. Vermeulen, 1906, fish trawl.
Paratype ZMA 06721, Mauritania, off Banc d’Arguin, 19.0667°N; 16.4167°W, dredged at 12–18 m, coll. R.W.M. Van Soest, R.V.’Tyro’ Mauritania II Expedition stat. 049, 11 June 1988.
Additional (non-type) material ZMA Por. 21697, Aegean Sea, fragment of specimen nr. 101 of E. Voultsiadou’s collections.
(
Skeleton: a confused mass of calthrops with a crust of microscleres at the surface.
Spicules: calthrops (no dichocalthrops) and sanidasters.
Calthrops (
Sanidasters (
Named after the locality of the holotype.
Consolidating coarse limestone fragments on rubble bottoms.
Senegal, Mauritania, possibly (Eastern) Mediterranean.
The new species was formerly identified as
The ZMA collection contains a tiny fragment of a specimen from the Aegean Sea donated by E. Voultsiadou (ZMA 21697) which has calthrops (
The record of
Holotype ZMA Por. 07521b, Cape Verde Islands, São Nicolau, Branco, 16.6667°N; 24.7167°W, dredged from 98 m, coll. R.W.M. Van Soest, HMS ‘Tydemann’ CANCAP VII Expedition stat. 156, 5 September 1986.
(
Skeleton: a confused mass of dichocalthrops with a thin crust of sanidasters.
Spicules: dichocalthrops (no calthrops), sanidasters.
Dichocalthrops (
Sanidasters relatively small, little variation in spination and ornamentation, 11–
Named after the type locality.
Deeper water, consolidating coarse sediment.
So far with certainty known only from the type locality. Possibly, Topsent’s 1928 record of
Although the specimen was originally identified with
BMNH 1907.2.1.5a, two slides from type, R.N. 167, Gulf of Mannaar, Sri Lanka.
(partially from Dendy, 1905). Encrusting extensively over calcareous algae and filling crevices, 4.5×3.3 cm in lateral expansion. Surface smooth, no apertures. Colour pale grey (alcohol). Consistency tough, fleshy. Dendy mentions the presence of large cells with inclusions (cystencytes) of 60 µm.
The two slides contain thin cross sections of the peripheral regions.
Skeleton (
Spicules: megascleres predominantly dichocalthrops, but also normal calthrops or calthrops with one or two bifid cladi, sanidasters.
Calthrops (
Dichocalthrops (
Sanidasters (
Encrusting calcareous algae at depths of 18–63 m.
Sri Lanka, ?Madagascar.
The spicule complement reminds of
It is likely that the record of
None. Holotype specimen, collected by R. Evans from Great Redang Island, Malaysia, could not be found in BMNH (2009).
(from Sollas, 1902). Sponge encrusting a dead coral fragment, 50×5×1–2 mm, surface smooth, shining; no oscules. Colour pink.
Spicules: Megascleres dominated by small dichocalthrops, but there are also small calthrops.
Calthrops have cladi of 60–70 × 3 µm (computed cladomes 90–115 µm diameter).
Dichocalthrops with protocladi 50–60 × 10 µm, deuterocladi 30 µm and rabdome 80 µm (computed cladomes 160–180 µm in diameter).
Sanidasters long and thin, with oxea-like shape: 15–20 × 1 µm.
Intertidal, occurring on dead coral fragments.
So far only known from Great Redang Island, one of the coral islands of the state of Trengganu at approx. 5°N on the E coast of Malaysia.
By the small size of the dichocalthrops this species appears similar to
None. The type material collected by Michaelsen & Hartmeyer from Shark Bay, West Australia could not be found in ZMB (listed as a slide ZMB 4492 in
(from Hentschel, 1909). Endolithic within corals, filling tubular cavities of 1–2 mm diameter which connect to the outside in only a few places. Colour (in alcohol) a deep brown. The skeleton consists of dichocalthrops and sanidasters.
Dichocalthrops relatively small, protocladi 20–28 µm, deuterocladi 50–92 µm, rhabd 86–105 × 12–18 µm (computed cladome diameter 130–230 µm).
Sanidasters 13–21 × 1.5 µm. Spines are described as strongly developed and irregular, but only tiny drawings of these spicules are provided, which only show elongated rhabd shapes.
An endolithic species from shallow coral reefs (0.5–3.5 m).
West Australia, Shark Bay, shallow water.
In spicule size this species appears close to
The only other species of
Holotype ZMUC unnumbered, Fiji Islands, Vitu Levu, Suva Harbour, coll. T. Wolff, 18 May 1965.
According to
Skeleton: unknown but presumably confused, with an ectosomal cover of microscleres.
Spicules: calthrops and dichocalthrops megascleres in approximately equal quantities, sanidasters.
Calthrops (
Dichocalthrops (
Sanidasters (
Named after its type locality.
Shallow-water, in crevices in dead corals.
Known only from the Fijian type locality.
The distinctive feature of this new species, apart from details of spicule sizes, is the characteristic shape of the larger dichotriaenes with very short protocladi and long conical deuterocladi (represented most clearly in the dichocalthops at the right of
The species closest to
Other species with a complement of both calthrops and dichocalthrops,
Holotype USNM 43170, Galapagos Islands, Albemarle Island, 0–3 m, 0.25°S; 91.4833°W, coll. Anton Bruun Cruise 16–66139, 25 May 1966 (slides and fragments of the holotype are present in MHNG, nr. 18963). Paratype ZMA Por. 11215, three fragments from the same locality.
(from
Skeleton: thick ectosomal crust of microscleres. Choanosomal skeleton confused mass of megascleres.
Spicules: calthrops, dichocalthrops, sanidasters. Some spicule types are remeasured to provide additional data and to correct a remarkable error in the original description.
Calthrops robust, variable in size, cladi 344–
Dichocalthrops, or short-shafted dichotriaenes, small, cladome 156–
Sanidasters: short and slim, with blunt apices. Remarkably, they were quoted to be 15–74 × 2–8 µm in size, but this is an obvious error. Remeasured they appear to be highly uniform in size and shape: 9–
Shallow water.
Only known from the Galapagos Islands.
By its large calthrops this species stands out among
Holotype HBOI 23–VIII–85–1–49, (HBOM 003.00040), with schizoholotype fragment ZMA Por. 07782, Bahamas, New Providence Island, NW of Goulding Cay, 25.025°N; 77.583°W, 210 m, coll. K. Rinehart in the Johnson SeaLink submersible, 23 August 1985, don. S.A. Pomponi and M.C. Díaz.
Slippery smooth encrustation on dead coral (
Skeleton: a confused mass of short-shafted triaenes and sanidasters.
Spicules: calthrops-like short-shafted triaenes, dichocalthrops and sanidasters.
Calthrops (
Regular symmetrical dichocalthrops (
Sanidasters (
Named for its type locality.
Collected by submersible at 210 m depth.
Known only from the Bahamian type locality.
This is a
None.
MNRJ 8559, Belmonte Islet Wall, São Pedro e São Paulo Archipelago, Brazil, 0.9167°N; 29.35°W, 30 m depth, coll. F. Moraes, 15 July 2004.
(from
Spicules. Calthrops 42–212 × 7.5–25 µm. Sanidasters 10–15 × 1 µm.
Vertical walls and overhangs, 4–30 m.
Offshore archipelagoes to the NE of Brazil.
The red colour is shared with
None.
USNM 21438, United States, California, Laguna Beach, coll. M.W. De Laubenfels, 26 March 1929.
(from De Laubenfels, 1932). Consolidating sandy substrate, shape ‘amorphous’, colour ‘drab’, surface smooth, consistency slimy. Size of parts containing the characteristic spiculation 0.2–2 mm. Skeleton of interior ‘packed’ with megascleres and microscleres.
Spicules: calthrops and sanidasters.
Calthrops, small, often reduced to ‘tripods’, cladi 25–
Sanidasters variable in spination, often concentrated in two areas of the microsclere to approach an amphiaster condition, but many are irregular, endings blunt, 8–12 µm.
Intertidal.
California.
None.
QMG329976 (SBD513022), south of Rock Cod Shoal, off Gladstone, Great Barrier Reef, 23.725°S; 151.6647°E, 34 m depth, epibenthic sled, coll. FRV ‘Lady Basten’, 20 September 2004.
(from
Spicules: three-claded calthrops and sanidasters.
Calthrops small, divisible in two size classes with means of approximately 25 and 72 µm. Only 20% of the 163 specimens recorded possessed calthrops, usually in high densities, in the remaining 80 % these spicules were lacking. Sanidasters universally present in all specimens, displaying a wide variation in length and width, 10–20 × 1–2.5 µm, densely spined with relatively short spines up to 1 µm.
Sandy bottoms between 16 and 86 m depth.
Great Barrier Reef, occurring over the entire range.
As
The only other Australian record of the genus,
None.
(from
Spicules: calthrops, sanidasters, ?strongyles.
Calthrops with irregular cladi, 50–250 × 15–25 µm.
Sanidasters short and fat, 12.5 × 2.5 µm.
Strongyles, probably foreign, long and thin 375–400 × 3–5 µm.
Shallow reefs.
Known only from the Tuléar area, Madagascar.
Live colour and the details of the spicules make it likely that this is an undescribed species. The strongyles are considered foreign as the authors themselves suggested.
ZMA Por. 08985, Netherlands Antilles, Bonaire, W coast, near Kralendijk, in reef caves, 12–43 m, 12.15°N; 68.278°W, 1984, coll. D.R. Kobluk.
Insinuating in coral crevices, in reef caves between 10 and 40 m depth. The available sponge material has been ‘used up’ entirely for the initial slides from which the genus identification (as
Calthrops regular, with four equal cladi, which may occasionally be curved; endings occasionally bifid or irregularly terminally knobbed, but no true dichotriaene nor triactine modifications are present; sizes highly variable, cladi 39–
Sanidasters, thin, 12–
In reef caves at the deeper parts of fringing reefs.
.
1 | Live colour yellow turning to dark brown or black in alcohol |
|
– | Live colour black | 2 |
– | Colour in alcohol beige or orange brown, compressed sanidasters ovoid, not euaster-like | 3 |
2 | Sanidasters very long up to 42 × 10 µm | |
– | Sanidasters long and thin, up to 33 × 6 µm | |
– | Sanidasters relatively short and fat, up to 27.5 × 7.5 µm |
|
3 | Insinuating habit, exclusively calthrops megascleres, microscleres include peculiar microspined ovoid rhabds | |
– | Encrusting habit, exclusively dichocalthrops megascleres, microscleres two sanidaster shapes, long-and-thin and fat-compressed |
Holotype, ZMB 287, Eritrea, Dahlak Archipelago, near Perim Island, Red Sea, 16.75°N; 40.05°E, 52 m, coll. Siemens.
Black mass (in alcohol) of 3×3×0.5 cm (
Skeleton: a confused, dense mass of calthrops with a cover of microscleres at the surface. A few oxeas of widely different sizes were noted by Keller, but these were limited between approx. 100–120 × 2.5 µm of a typical haplosclerid type, in the slide examined by us.
Spicules: calthrops, sanidasters, compressed sanidasters (pseudasters).
Calthrops (
Sanidasters (
Compressed sanidasters (pseudasters) (
Deeper water, beyond the reefs
Known only from the Southern Red Sea.
Holotype RMNH 4256, Indonesia, Kalimantan, Berau region, Derawan Islands, depth 10 m, coll. N.J. De Voogd, field nr. BER105/140808/055, SCUBA, 14 August 2008.
Blackish grey lobate mass (
Skeleton: difficult to study in the preserved condition due to the intense black colour, but structure is dense and unorganized, with a dense cover of microscleres at the surface.
Spicules: calthrops, sanidasters and compressed spheraster-like microscleres (pseudasters); a few oxeas are present, but these belong to the encrusting
Calthrops (
Sanidasters (
Compressed sanidasters (pseudasters) (
Named after the Berau region, East Kalimantan.
Shallow open reef localities.
So far known only from the type locality E of Kalimantan.
Despite the long distance between the locality of
Holotype ZMA Por. 17646 (voucher fragment of a specimen used in natural products research) Japan, near Anami-oshima Island, 157–161 m, 28.873°N; 129.5532°E, coll. N. Fusetani, dredge, field nr. S01–111, 7 November 2003.
Paratype ZMA Por. 19889 (voucher fragment of a specimen used in natural products research), Japan, Kagoshima Prefecture, Ooshima-shinsone, 150 m, 28.8667°N; 122.55°E, coll. Y. Nakao, dredge, field nr. S01–118, 16 July 2001.
Holotype (
Skeleton: at the surface a crust of microscleres, overlying a dense confused mass of calthrops.
Spicules: calthrops, sanidasters, compressed sanidasters (pseudasters).
Calthrops of holotype (
Sanidasters of holotype (
Compressed sanidasters of the holotype (
Named after the Japanese type locality.
Dredged from deeper water, at approx. 150 m.
Both specimens came from nearby locations, so the distribution so far appears limited to offshore Japan.
It is with some hesitation that these two Japanese sponges are assigned to a new species, rather than to
Spicule sizes of specimens similar to
Author | calthrops cladi | sanidaster | compressed sanidaster | locality |
---|---|---|---|---|
|
||||
|
not given | not given | ½ sanidaster size | Red Sea |
|
200×25 | 25 | 5–10 | Red Sea |
Remeasured here | 45–275×5–35 | 15–27.5×2.5–7.5 | 7.5–12.5 | Red Sea |
RMNH 4256 | 25–280×5–35 | 19–42×1–10 | 8–15 | Indonesia |
ZMA Por. 17646 | 105–223×14–28 | 27–31×2.5–4.5 | 7.5–15 | Japan |
ZMA Por. 19889 | 33–252×5–37 | 19–33×2–6 | 7–14 | Japan |
ZMA Por. 10100, Netherlands Antilles, Curaçao, Boca Sami, 20 m, 12.142°N; 69.007°W, coll. P. Willemsen nr. C92 A20, July 1992.
ZMA Por. 14080, Netherlands Antilles, Curaçao, near Carmabi, in reef cave, 10 –20 m, 12.124°N; 68.974°W, coll. I. Wunsch nr. 84, January 1999.
ZMA Por. 21692 (ex MNRJ 6678), Brazil, Das Rocas Atoll, Fenda, 3.8584°S; 33.8057°W, coll. E. Hajdu, M.V. de Oliveira & U.S. Pinheiro.
The larger specimen (10100) is a thick liver-like mass (
Skeleton: the ectosome has a dense layer of sanidasters and ‘spherasters’ carried by a similarly dense mass of calthrops. In the choanosome the calthrops and sanidasters are less densely and confusedly distributed. There is no zonation or other structural arrangement of the skeleton.
Spicules: calthrops, sanidasters and compressed spheraster-like sanidasters (pseudasters).
Calthrops (
Sanidasters (
Compressed sanidasters (
Open substrate in deeper parts of the reef, more shallow in caves.
Widespread Central West Atlantic: Puerto Rico, Southern Caribbean, Northeastern Brazil.
The Curaçao material appears indistinguishable from the Puerto Rico type material (MSNG 47681, La Parguera, Puerto Rico, 20–25 m) in overall morphology and spicules. Pulitzer’s material measures 7×5×1.5 cm, turning from light yellow to dark brown in alcohol. Consistency was described as fleshy, firm, compressible. Cladi of the calthrops were given as 70–180 µm (60–174 in our material), sanidasters (called microrhabds by Pulitzer) were given as 8–26 × 3–6 µm (12–26 × 1–4 µm in our material). Compressed sanidaster size was not given by Pulitzer but from his drawings can be estimated to be about 7–8 µm (5–9 µm in our material).
Through the courtesy of Prof. Hajdu we received a fragment (
Spicule data for specimens identification as
|
|
|
|
|
---|---|---|---|---|
|
70–180×× | 8–26×3–6 | 7–8 | Puerto Rico |
ZMA 10100 | 60–174 ×9–21 | 12–26×1–4 | 5–9 | Curaçao |
ZMA 14080 | 80–130×15 | 18–21×3–4 | 7–9 | Curaçao |
ZMA 21692 (MNRJ 6678) | 33–123×4–11 | 10–19×0.5–2 | 5–9 | Oceanic islands Brazil |
Holotype ZMA Por. 08984, Netherlands Antilles, Aruba, Lagoon Boekoeti, underneath coral rubble, 0–1 m, coll. P. Wagenaar Hummelinck, nr. 1004, 29 December 1948.
Small soft patches of a brown-red colour insinuating in three coral fragments (
Skeleton: Confused mass of calthrops and microscleres, no explicit structure.
Spicules: Calthrops, sanidasters and microrhabds.
Calthrops (
Sanidasters (
Microrhabds (
Named after the holotype locality.
Shallow-water, under coral rubble.
So far known only from the island of Aruba, Southern Caribbean.
The microrhabds are unique among species of the genus
Holotype ZMA Por. 02220, Indonesia, Papua, 32 m, 1.7083°S; 130.7916°E, coll. Siboga Exped. Stat. 164, dredge, 20 August 1899.
Thin leathery-rubbery encrustation (
Skeleton: an ectosomal crust of microscleres overlying a mixture of microscleres and dichocalthrops embedded in largely organic choanosome with relatively low spicule density.
Spicules: dichocalthrops, sanidasters and compressed fat sanidasters.
Dichocalthrops (
Sanidasters in two categories, long-and-thin (
Named after the Dutch naval vessel HMS ‘Siboga’ which was instrumental to the 1899–1900 collection of a rich deep sea fauna from South East Asian waters.
Dredged from a sandy bottom with small limestone rubble and shells.
Known only from the type locality in Eastern Indonesia.
The present specimen was identified by Maurice Burton (who provisionally identified a large part of the Siboga sponge collection) as
None.
(from Lebwohl, 1914). Encrusting a lithistid (
Spicules: calthrops, microrhabds, metasters.
Calthrops variable in number of cladi, often with bifid endings, but no real dichocalthrops, quite variable in size: 40–1250 × 5–165 µm.
Microrhabds ovoid, occasionally globular, rugose or microspined, 11.5–17.5 × 5.5–8.5 µm.
Metasters, microspined, 14–26 µm, with 4–18 rays of 5–20 µm length.
Deep water, 214 m.
Japan.
This is a dubious description as it was partly overgrown by a
None.
(from Burton, 1926). Encrusting on
Deeper water.
Natal coast, South Africa.
This is a dubious
Comments:
The genus
The junior synonymy proposed for
The newly defined genus
.
1 | Microscleres include only ‘normal’ asters (oxyasters and strongylasters) |
|
– | Microscleres include next to ‘normal’ asters also highly silicified globular or irregular shapes | 2 |
2 | Microscleres include ‘ataxasters’ (irregular branched or ovoid microspined forms) |
|
– | Microscleres include regular tuberculated globular asters |
|
Key to the subgenera of
.
1 | Dichocalthrops present | 2 |
– | No dichocalthrops | 3 |
2 | Cladi of calthrops up to 800 µm |
|
– | Cladi of the calthrops only 210 µm | |
3 | Larger calthrops predominantly reduced to ‘tripods’ (three-claded megascleres); smaller microscleres deformed to form highly silicified spherical bodies spined all over |
|
– | Larger calthrops variable, not predominantly ‘tripods’; microscleres recognizably spherasters and oxyasters | 4 |
4 | Calthrops extremely variable in shape, many with reduced number of cladi | 5 |
– | Calthrops regular, but may have an extra cladus | 6 |
5 | Two categories of oxyasters; smallest oxyasters smooth |
|
– | Single oxyaster category, all microspined |
|
6 | Oxyasters in two size categories | |
– | Oxyasters in a single size category |
|
None. Type material could not be found in the Natural History Museum (query in 2009).
(from
Spicules: calthrops, globular spherasters and ‘normal’ oxyspherasters.
Calthrops, of widely different sizes, according to Sollas divisible into larger (cladi dimensions 800 × 90 µm) and smaller (150 × 20 µm).
Spheroxyasters, considered a young stage of the tuberculated asters by Sollas, but identified as a proper separate aster-type by Topsent; no size given by either authors but according to Topsent the same size as the truncated asters, so 12–24 µm in diameter.
Globular spherasters with tuberculated rays, variable, often irregular, in a wide size range, possibly divisible in two size categories, 24 µm and 12 µm.
Deep water, on gravel/sand bottom. Sollas provides no depth data, Topsent’s material came from 454 –861 m.
Cape Verde Islands (off Porto Praya, Santiago Island), Azores (38–39°N 30°W).
This species differs from the closely similar
Spicule data for
Author | calthrops cladi | oxeas | tuberculated sphaerasters | oxyasters | dicho- calthrops |
---|---|---|---|---|---|
|
|||||
150–800×20–90 | not recorded | 12–24 | yes | no | |
|
|||||
700×85 | 736×10 | 25 | not recorded | yes | |
yes, large variation | not recorded | 20 | 12–15 | yes | |
present paper (Azores) | 102–705× 11–128 | broken, few | 7–28 | 13–18 | yes |
|
|||||
present paper (Azores) | 61–679× 4–96 | 800–1450 9–13 | (1) 7–30 (2) 7–36 | (1) 18–24 (2) 11–18 | no |
|
|||||
550×75 | long × 15 | 20–23 | 12–15 | no | |
present paper MNHN DT 754 | 32–366× 5–72 | 2000×12 | 9–24 | (1) 23–27 (2) 9–12 | no |
60–800× 5–100 | long × 12 | 15–22 | 12 | no | |
40–600× 5–70 | broken × 5–12 | 10–18 | 10–18 | no | |
50–495× 3.6–50 | 2000×15 | 7–17.6 | 7.2–17.6 | no | |
|
|||||
34–600 | 2000×15 | 5–22 | 8–17 | no | |
present paper (Indonesia) | 44–587× 5–101 | not present | (1) 18–27 (2) 6–10 | (1) 15–21 (2) 8–10 | no |
|
180–220 | 750×20 | 10–12 | not recorded | yes |
ZMA Por. 21666, EMEPC/G3–D03A–Ma012, Azores, Terceira Island, 38.4265°N; 26.8206°W, 1201 m, coll. J. Xavier, 18 May 2007.
Specimens provided by Dr Joana Xavier, as yet unregistered: EMEPC/G3–D01–Ma005, Azores, São Jorge Island, 38.48°N; 27.798°W, 1222 m, 17 May 2007; EMEPC/G3–D02–Ma006, Azores, Terceira Island, 38.5530°N; 26.7083°W, 744 m, 18 May 2007; EMEPC/G3–D33A–Ma011a, S of Azores, Cruiser Seamount, 32.2570°N; 27.553°W, 643 m, 4 June 2007; EMEPC/G3/08.30, S of Azores, Atlantis Seamount, 33.916°N; 30.171°W, 1132 m, 30 October 2008.
Flat mounds (
Skeleton: with a dense crust of microscleres carried by triaene megascleres, with scattered bundles of oxeas, which appear mostly broken and are sometimes entirely absent.
Spicules: calthrops, dichocalthrops, (broken) oxeas, tuberculated spherasters, oxyasters.
Calthrops (
Dichocalthrops (
Oxeas (visible in
Oxyasters (
Tuberculated spherasters (
On rocks in deep water, 599–1222 m.
Off Cabo São Vicente, Portugal; Azores.
We believe that the two species
Topsent’s (1897) variety of
Specimen provided by Dr. Joana Xavier, as yet unregistered: EMEPC/G3/08.30, S of Azores, Atlantis Seamount, 33.916°N; 30.171°W, 1132 m, 30 October 2008.
Not examined. Dr. P. Cárdenas (
White, massive lump (
Skeleton: largely confused interiorly, with a densely crowded outer layer of microscleres.
Spicules: calthrops, two types of oxyasters, tuberculated asters, lumpy asters; some broken oxeas.
Calthrops (
Oxeas (
Oxyasters, occurring in two distinct types: small oxyasters (
Tuberculated asters (
Lumpy, irregular aster-derived silica bodies (
Deep water, 450–1132 m.
W of Flores and seamounts S of the Azores.
Topsent’s variety
Lectotype (designation herein), MNHN DT 753, Coast of Algiers, Expédition Scientifique de l’Algérie, nr. 66, 1842. Paralectotype MNHN DT 754, from same locality.
(partly from Topsent, 1938). Two specimens are present in the Schmidt collection of the Muséum National d’Histoire Naturelle, Paris, the largest (MNHN DT 753) of which is 4.5 cm in widest size and 2–2.5 cm high, here chosen as the lectotype. The smaller specimen (MNHN DT 754), here designated paralectotype, size 1×3 cm (now apparently reduced to 1×1.5 cm) was examined by us and we made SEM photos of the spicules. Both specimens have a rough surface (
Spicules: Calthrops, tuberculated spherasters, oxyasters.
Calthrops (
The megasclere complement also comprised thin oxeas of 15 µm thickness (
Asters (
Lightly spined oxyasters (
Small, smooth oxyasters (
Tuberculated asters with ornamented rays (
Deep water, and in caves in more shallow water, depth occurrence at least 13–250 m.
Off the coast of Algeria, no further data; Cassidaigne, Marseille region; Sicily-Tunis region; Alboran Sea, 39°N –3°W; Rhodos, Aegean Sea, 36°N; 28°E.
None.
MSNG 47158, off Camogli, N Italy.
(from
Spicules: calthrops, oxeas, strongylasters to spherasters, oxyasters.
Calthrops, extremely variable in shape, with many reductions and malformations, size variation also considerable, 34–600 µm (no thickness given).
Oxeas, with extremely elongated, flexuous points, all broken in the slide, but at least 2000 × 15 µm.
Oxyasters, without centre, with two to six microspined rays, 8–17 µm.
Strongylasters to spherasters, variable in shape, irregular with tuberculated rays or lobate, 5–22 µm.
Trawled from 60 m depth.
Known only from the type locality off the coast of Genoa, 44°N; 9°E.
This species is similar to
Pulitzer (l.c.) suggests that
Holotype ZMA Por. 11376, Indonesia, E of Komodo, 8.4867°S; 119.6167°E, depth 138 m, dredge, coll. R.W.M. Van Soest, Snellius II Exped. Stat. 095, 19 September 1984.
Lobate mass (
Skeleton: a dense crust of asterose microscleres at the surface covers a dense mass of calthrops. Asters are also crowded in the choanosome. The skeleton is densely confused and there are few canals or cavities; some broken monaxone spicules were present.
Spicules: calthrops, globular tuberculate asters, oxyspherasters, oxeas?.
Calthrops (
Broken monaxonic spicules (oxeas?) measured up to 700 × 10 µm.
Oxyasters (
Globular tuberculated spherasters (
The name of the new species acknowledges all the help we received from Joana B.R.T. Xavier during her stay in Amsterdam, and the generous donation of several sponge fragments important for this study.
Dredged from hard bottom at 138 m.
Known only from the type locality, E of the island of Komodo, Nusa Tengara (southern island chain bordering the Banda Sea).
This is a clear
No matching descriptions are found in the Indo–West Pacific region, but
None. According to
(from
Shallow reefs.
Ambon Bay, Indonesia.
It is likely that the Ambon material belongs to an undescribed species, as the length of the cladi of the calthrops and the diameter of the tuberculated asters are much smaller than in the type of
.
1 | Oxyasters up to 24 µm, ataxasters predominantly pear-shaped | |
– | Oxyasters less than 12 µm, ataxasters irregularly branched | 2 |
2 | Ataxasters up to 48 × 34 µm, sponge plate-shaped |
|
– | Ataxasters up to 42 × 16 µm, sponge lobate |
|
Holotype MCZ 6384, Blake Exped. 1878–79, Grenada, 160 m. Further type material: schizoholotype fragment MZUS P0095 (not examined) and two type slides in ZMB, nr. 6870 (not examined).
Thick plate (
Skeleton: a confused mass of calthrops with at the periphery a dense mass of ataxasters, which are also strewn in the interior.
Spicules: calthrops, spheroxyasters, ataxasters.
Calthrops (
Spheroxyasters (
Ataxasters, of many different shapes and sizes (
Deep water, 160 m.
Only known from the type locality, off Grenada.
The type material was assigned by
Holotype MNHN DCL 2805; paratypes DCL 2895 (4 small sponges), both from New Caledonia, 22.8°S; 167.15°E, depth 355–360 m.
Massive with rounded flattened lobate outgrowths (
Skeleton: confused, with a crust of microscleres carried by the triaene megascleres. A few broken oxea-like spicules were present in the slides made from the largest paratype.
Spicules (
Calthrops (
Spheroxyasters (
Ataxasters (
Deep water, 355 m.
Known only from off New Caledonia.
The New Caledonian species differs from the Caribbean
Holotype HBOI nr. 12–XI–86–1–14, with fragment (schizoholotype) ZMA Por. 07726, Ecuador, Galapagos Archipelago, NE coast of Santa Cruz, Rocas Gordon, 0.546°S; 90.116°W, depth 506 m, coll. K. Rinehart, Johnson SeaLink submersible, 12 November 1996, don. S.A. Pomponi & M.C. Diaz.
Type material available to us has a flattened (
Skeleton: an irregular mass of large triaenes covered at the periphery by a thick layer of ataxasters / microrhabds, which are also strewn in the interior. Loose oxeas of widely different sizes present in moderate quantities, considered foreign.
Spicules: calthrops, spheroxyasters, ataxasters.
Calthrops (
Spheroxyasters (
Ataxasters (
From the Greek pyros = pear and fero= carry or bear, referring to the possession of the pear-shaped microscleres.
Deep water, 506 m.
NE of Santa Cruz Island, Galapagos, East Pacific.
The species is assigned to
.
1 | Calthrops cladi small and thin, less than 195 × 10 µm | |
– | Larger and thicker calthrops | 2 |
2 | Oxyasters up to 70 µm diameter |
|
– | Oxyasters only up to 50 µm diameter | 3 |
3 | Auxiliary oxea megascleres present |
|
– | No oxeas |
|
Type fragment of LMJG 15352 from Sebenico, Northern Adriatic (slide in ZMA). There are further fragments, not examined, BMNH 1867.7.26.14, 45, 104, BMNH 1910.1.1.863, MZUS P0005, and slides BMNH 1868.3.2.5 and ZMB 6563).
From
Skeleton: densely spiculous at the surface, organic with few spicules in the interior. No definite skeletal structure.
Spicules: calthrops, euasters.
Calthrops (
Microscleres euasters in two categories (although the LMJG fragment available to us only contained a single one: strongylasters 16–
Fairly shallow water, from intertidal caves to 20 m.
Adriatic; Marseille; Canary Islands; NW Aegean Sea, 39°N; 25°E (Algeria is given by previous authors, but this is probably based on a misunderstanding of Schmidt’s text).
The species is the type of the preoccupied genus
None. Apparently the material that is left consists only of slides, of which the holotype is in the Natural History Museum, London (BMNH 1971.7.23.1b).
(from
Spicules: calthrops, oxeas, strongylasters (called ‘chiasters’), oxyasters.
Calthrops in a large size variation and many showing abnormal or assymmetrical cladi (the latter distinguished as a separate category by Pulitzer), cladi 85–220 × 7–17 µm.
Oxeas straight or curved, in a large size range, 100–1200 × 7.5 µm.
Strongylasters, 8–12 rays, variable, 9–16 µm.
Oxyasters, six thin rays, 40–50 µm.
Collected at 12 m.
Mediterranean (Naples region).
The oxeas are explicitly stated as part of the spicule complement, but were not represented in the spicule drawings. It remains doubtful whether they are proper.
None.
(from
Skeleton: no data.
Spicules: calthrops, strongylasters, oxyasters.
Calthrops, occasionally with five or six cladi, 120–210 × 26–38 µm; “occasionally with dicho-modifications”; the 1968 illustration (Fig. 29) shows at least one proper short-shafted dichocalthrops.
Strongylasters, rather irregular in shape, with stunted rays, but not with a thickly silicified centre, 11 µm.
Oxyasters, 50–70 µm (5–7 rays).
Sublittoral fringe.
Rangitoto Island, northern New Zealand.
The drawings and photos of the asters are limited to the rather irregular strongylasters, whereas no illustrations are available of the oxyasters. It is assumed these were regular and unremarkable.
ZMA Por. 10525, Seychelles, Mahé, Cap Maçons & Anse de Forbans, 4.7667°S; 55.5167°E, 0–6 m, NIOP Expedition stat. 612, coll. R.W.M. Van Soest, nr. 612/20, 12 December 1992.
Cartilaginous crust on a piece of dead coral, surface smooth, size 2×1×0.5 cm. Greyish black alive, pale brown in alcohol.
Skeleton: with low spicular density, predominantly consisting of calthrops.
Spicules: Calthrops, euasters in two categories.
Calthrops (
Oxyspherasters (
Small oxyasters (
Encrusting dead corals in shallow water.
Known only from a single locality south of Victoria, Mahé.
We refrain from naming this material due to the rarity of its microscleres. By its small sized calthrops this is a distinct species, as no matching descriptions appear in literature dealing with
None. Holotype MSNG 48290
Branching mass of 5×3 cm. Confused skeleton in which giant oxeas (2700 × 9 µm) predominate. Next to this there are large dichotriaenes with cladomes of 400–550 µm wide and similar sized rhabdomes of 400–650 × 35–45 µm. Spherasters of 8–14 µm comprise the microscleres.
Deep water, 120 m.
North Kenya Banks, E Africa.
This is in all probability not a
From the above presented data a close relationship of
Assuming that the similarities will be found to be homologous and that the two considered groups will be found to be closely related, family assignment to one of the currently recognized families is not unequivocal.
Including the three unnamed species described above, there appear to be at least seven records of
–
–
–
–
–
–
–
On the other hand, there are at least six species suspected to be junior synonyms:
–
–
–
–
–
A further two records (
Previous analysis of generic distributions of
Distribution of species of the genus
Distribution of species of the genus
Dr Joana Xavier (CIBIO, Azores) contributed by allowing us to study specimens of