Prosadenoporus agricola ( Willemoes-Suhm, 1874 ) Maslakova & Norenburg, 2008

Maslakova, Svetlana A. & Norenburg, Jon L., 2008, Revision of the smiling worms, genera Prosadenoporus Bürger, 1890 and Pantinonemertes Moore and Gibson, 1981 and description of a new species Prosadenoporus floridensis sp. nov. (Prosorhochmidae; Hoplonemertea; Nemertea) from Florida and Belize, Journal of Natural History 42 (25 - 26), pp. 1689-1727 : 1697-1702

Prosadenoporus agricola ( Willemoes-Suhm, 1874) , new combination

( Figure 10F ; Tables 2 and 3)

Tetrastemma agricola ( Willemoes-Suhm 1874; Moseley 1879; Hubrecht 1887; Verrill 1902).

Neonemertes agricola ( Girard 1893; Joubin 1894; Friedrich 1955).

Geonemertes agricola ( Bürger 1895, 1897 –1907, 1904; Coe 1904, 1905, 1929, 1939, 1940; Crozier 1917; Pantin 1947, 1961, 1969; Gibson 1972; Moore and Moore 1972; Moore 1973, 1975a, 1975b).

Pantinonemertes agricola ( Moore and Gibson 1981; Gibson et al. 1982; Gibson 1982b; Gibson, 1990; Sundberg 1989; Gibson and Sundberg 1992; 2001; Moore et al. 2001; Sun 2001).

Etymology

The name reflects terrestrial habitat of this species ( agricola 5farmer, L).

Type material

Types were never designated. Original material almost certainly lost.

Material examined

Prosadenoporus agricola ( Willemoes-Suhm, 1874) comb. nov. Series of sections of one mature and one juvenile specimen are held at the University Museum of Zoology, Cambridge, UK (coll. Boden, B. 1951, Bermuda).

Diagnosis

Prosadenoporus agricola comb. nov. differs from most other Prosadenoporus species in being hermaphroditic and viviparous ( Table 3). It differs from Prosadenoporus arenarius , the only other hermaphroditic congener, whose life history is unknown, by having fewer proboscis nerves 12–15 (compared with 15–17) and possessing neurochord cells. Central stylet (S) 80–95 mm long, basis (B) rounded, pear-shaped or truncated 75–90 mm long, two accessory stylet pouches. It is not possible to determine mean S:B ratio because only ranges of stylet and basis lengths are available. Data at hand are insufficient to determine whether these metrics are significantly different from those of other species.

Habitat and distribution

Along the shores of mangrove swamps, under stones and logs on moist, silty soil or inside burrows of earthworms above high-water mark and, during wet season, on adjacent hillsides. Associated with a heteronemertean Lineus sp., earthworms, nematodes and, sometimes, insects. Smaller individuals also occur below the highwater mark. Found in Hungary Bay, Bayley’s Bay, Walsingham Bay, on shores of Castle Harbor, near the ‘‘Causeway’’, at Hamilton Harbor and other localities on Bermuda Island. Once common on Bermuda ( Coe 1904, p. 532), this species had last been collected in 1966 during an extensive search of the islands by Dr B. Boden, which yielded very few specimens, all obtained from a remote place. More recent commercial development has left very little suitable habitat and the species has apparently not been found since ( Moore et al. 2001; W. Sterrer, personal communication to SAM).

Remarks

Coe’s (1904) anatomical account of the species mentions 13–15 proboscis nerves. SAM observed 12 proboscis nerves in the available voucher specimen in the collection of University Museum of Zoology, Cambridge, UK.

Prosadenoporus arenarius Bürger, 1890

Prosadenoporus arenarius ( Bürger 1890, 1895, 1904; Moore and Gibson 1988).

Etymology

The species name means ‘‘living in sand’’ (L).

Type material

Type material almost certainly does not exist; not known from the most likely repositories – Museum für Naturkunde, Berlin; Zoologisches Institut, Universität Göttingen; Stazione Zoologica, Naples.

Material examined

Prosadenoporus arenarius Bürger, 1890 . Series of transverse sections (slides 1–3) of hermaphroditic specimen from Punnett’s collection ( Palau Bidan off Malay Peninsula) held at the University Museum of Zoology, Cambrige, UK.

Diagnosis

Prosadenoporus arenarius differs from all other Prosadenoporus species except Prosadenoporus agricola comb. nov. by being hermaphroditic ( Moore and Gibson 1988, p. 81) and from the Bermudan P. agricola by having 15–17 proboscis nerves (compared with 12–15) and by lacking neurochord cells ( Table 3). Characters of the proboscis armature are unknown.

Habitat and distribution

Habitat is unknown, except in as much as the name of the species implies. Found in Java Sea (Noordwachter Island off northwestern Sulawesi) and Palau Bidan (Kedah Province, off Malay Peninsula).

Remarks

As mentioned above, Bürger’s material is almost certainly lost. A tube with several specimens labelled ‘‘ Prosadenoporus sp.’’ from Punnett’s collection from Palau Bidan (Kedah Province, Malay Peninsula) was deposited at the University Museum of Zoology in Cambridge, UK. Moore and Gibson (1988) sectioned some of these specimens and re-described Prosadenoporus arenarius Bürger, 1890 , arguing that Punnett’s specimens fit the original (very brief) description of P. arenarius . However, some evidence suggests that Bürger’s P. arenarius from Indonesia and Punnet’s Prosadenoporus sp. from Malaysia might represent different species.

First, the specimen described by Bürger (1890) from the Noordwachter Island off Sulawesi had a broad dark-brown, mid-dorsal stripe on a grayish-green background, while Punnett’s specimens were uniformly dull brown ( Moore and Gibson 1988). Second, and more important, Bürger’s diagnosis of the genus mentions neurochord cells and neurochords. Moore and Gibson (1988) report about Punnett’s specimens that ‘‘few neurochord cells are present, located near the inner margins of the dorsal cerebral lobes and neurochords can be traced in the main nerve cords’’ (p. 80). However, their figure 10 ( Moore and Gibson 1988, p. 78, fig. 10) suggests that they mistake the type III neurons for neurochord cells, both defined by Bürger (1895). Our reinvestigation of Punnett’s material showed no traces of neurochord cells or neurochords. Finally, Bürger’s specimen had 12 proboscis nerves, while there are 17 proboscis nerves in Punnett’s material. Although we have little confidence that Bürger’s description and Punnett’s material belong to the same species, lack of the type or other informative material thwarts resolution of the question.

Moore and Gibson (1988, p. 82) provided an amended diagnosis of the genus Prosadenoporus . The main difference between Pantinonemertes Moore and Gibson, 1981 and Prosadenoporus Bürger, 1890 is the presence of a second cephalic vascular loop described for Prosadenoporus by Moore and Gibson (1988, p. 81, fig. 18). However, the two authors of the present paper, as well as Frank Crandall (National Museum of Natural History, Smithsonian Institution, personal communication), independently confirmed the presence of a single cephalic vascular loop upon reexamining Punnett’s material. This eliminates the main distinction between the two genera and provides grounds for synonymizing Prosadenoporus Bürger, 1890 and Pantinonemertes Moore and Gibson, 1981 .

Prosadenoporus enalios ( Moore and Gibson, 1981) , new combination ( Figures 7 A–C and 10E; Tables 2 and 3).

Pantinonemertes enalios ( Moore and Gibson 1981; Gibson 1982b, 1990; Gibson et al.

1982; Sundberg 1989; Gibson and Sundberg 1992; Sun 2001).

Etymology

The species name reflects the fully marine habitat of the species; ( enalios 5in the sea, G).

Type material

Prosadenoporus enalios ( Moore and Gibson, 1981) comb. nov. Sections of holotype W5900 and paratype W5899 are held at the Australian Museum, Sydney, Australia.

Material examined

Prosadenoporus enalios ( Moore and Gibson, 1981) comb. nov. Holotype W5900, paratype W5899. Additional sectioned specimen 1978-12-3, coll. by R. Gibson from Magnetic Island , Queensland, Australia held at the Natural History Museum, London, UK.

Diagnosis

Prosadenoporus enalios comb. nov. differs from P. fujianensis , P. winsori and P. spectaculum by paler colouration and the absence of neurochord cells. Additionally, it differs from P. winsori in lacking neurochords. It differs from P. agricola and P. arenarius by being gonochoric, and from P. mortoni and P. mooreae by lacking characteristic greenish striped colouration and by having a truncated basis of central stylet ( Figure 10E ). It differs from P. floridensis sp. nov. by the absence of neurochord cells and having 14–16 proboscis nerves (compared with 11–14). Number of accessory stylet pouches unknown. Central stylet (S) 110 mm long, basis (B) truncated 170 mm long, S:B ratio 0.65 ( Moore and Gibson 1981). Data at hand are insufficient to determine whether these metrics are significantly different from those of other species.

Habitat and distribution

In silty mud beneath rocks and coral boulders, mid- to lower-shore, intertidal. Nelly Bay and Picnic Bay on Magnetic Island, Queensland and on Pelorus Island, Palm Island Group, Great Barrier Reef, Australia.

Remarks

Original description mentions 14–15 proboscis nerves ( Moore and Gibson 1981). However, we observed 16 nerves in the holotype. Although it appears to have been abundant at the time of original description, SAM failed to re-collect any from the type locality and other localities nearby in March 2003 despite the successful recollection of P. mooreae from the same area.

Figures

Figure 7. Microscopic anatomy of Prosadenoporus species.(A–C) A series of transverse sections through the frontal organ of Prosadenoporus enalios comb. nov.; relative position of the lateral acidophilic regions (arrowheads) indicate a noticeable twist of the frontal organ canal; (D) a longitudinal sagittal section through frontal organ of Prosadenoporus mooreae comb. nov. (anterior to the right); (E–I) a series of slightly oblique transverse sections through the frontal organ of P. mooreae comb. nov.; lateral acidophilic regions of the frontal organ (arrowheads), note the twisting of the frontal organ canal; (J) transverse section through lateral nerve cord of Prosadenoporus winsori comb. nov. showing the tentative neurochord (arrowhead); (K) binucleate terminal cells of nephridia (flame cells) of P. winsori comb. nov. (arrowheads); (L) a longitudinal sagittal section through the frontal organ of P. winsori comb. nov. (anterior to the left). Notes: bgl, basophilic cephalic glands; cg, cerebral ganglia; fo, frontal organ; lnc, lateral nerve cord; rhd, rhynchodeum. Scales: (A–C), and (E–I), 50 mm; (D), 100 mm; (J), 30mm; (K), 20mm; (L), 200mm.

Figure 10. Central stylets: (A) Prosadenoporus mortoni comb. nov. (a sketch of stylet apparatus provided by Dr Shichun Sun); (B) Prosadenoporus mooreae comb. nov. (specimen from the type locality coll.by SAM); (C) Pantinonemertes californiensis (after Gibson et al. 1982, p.471, fig.3b); (D) Prosadenoporus fujianensis comb. nov. (redrawn from Sun 2001, p.205, fig.23); (E) Prosadenoporus enalios comb. nov. (redrawn from Moore and Gibson 1981, p.183, plate III, fig.c); (F) Prosadenoporus agricola comb. nov. (redrawn from Coe 1904. p.571. plate 23, fig.3); (G–H) Prosadenoporus winsori comb. nov. (specimen from the type locality coll. by SAM); (I) Prosadenoporus floridensis sp. nov. Note: Scales: (A, B, E, F), 100mm; (C, D, G, H, I), 500 mm.