Research Article |
Corresponding author: Christopher J. Glasby ( chris.glasby@nt.gov.au ) Academic editor: Robert Blakemore
© 2015 Joko Pamungkas, Christopher J. Glasby.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pamungkas J, Glasby CJ (2015) Taxonomy of reproductive Nereididae (Annelida) in multispecies swarms at Ambon Island, Indonesia. ZooKeys 520: 1-25. https://doi.org/10.3897/zookeys.520.9581
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Multispecies, or mass, spawning of different invertebrate species is well known for coral reef systems; however, incidences involving polychaetes are poorly documented. In this study we report on mass swarming, prior to spawning, of Nereididae at Ambon Island, Maluku, on three occasions: in 1866, inferred from an historical sample deposited in Naturalis, Leiden, and in March, 2009 and 2014, based on newly collected samples. The 2009 and 2014 events co-occurred with spawning of other polychaetes, known locally as wawo and including the widespread Indo-Pacific eunicid, Palola viridis (Gray in Stair). Ten species of reproductive Nereididae are described, including Composetia marmorata (Horst) new combination, formerly Ceratonereis marmorata; epitokous modifications are described for both sexes of each species including taxonomically important features such as body colour and number of pre-natatory chaetigers. Three distinct types of natatory region morphologies are recognized, which appear to characterise groups of genera. The ten new records brings to 13 the total number of nereidid species known to undergo mass swarming at Ambon Island; a key to the 13 species is provided. Species composition varies slightly between the three time periods: four species were common between all three periods, five species were in common between 1866 and 2014, and four species were in common between 1995 and 2009/14. Two species of Neanthes and one of Nereis are identified as potentially new and will be described in subsequent papers.
Systematics, heteronereid, epitoke, new species, Polychaeta, Maluku, Wallacia
Synchronised swarming of polychaetes at the surface of the sea for the purpose of breeding is perhaps best known among the family Eunicidae, in particular the Indo-Pacific species Palola viridis Gray in Stair, 1847. However, swarming is also common among Nereididae and Syllidae and occurs also in at least 15 other families (
In Ambon and surrounding islands of Maluku, mass swarming of polychaetes occurs every year from February to April. Locals refer to the worms as ‘wawo’ or, more popularly, as ‘laor’. The swarming has been known for a very long time, as occurring either in February and March (
The only previous taxonomic studies of Ambonese wawo are those of
In this study we describe the taxonomy of the nereidid component of wawo collected in 2009/14 and nereidids in Hoedt’s collection (1866), in particular the epitokous modifications of each sex and species. Because all samples contained a small fraction of spent worms (i.e., lacking coelomic gametes), we have assumed that spawning occurred about the time the swarming worms were collected. The species composition of the two collections, almost 150 years apart, is compared with that reported by
Sexually mature polychaetes (wawo) were collected during the swarming period on 14 March, 2009 and 18–19 March, 2014, from Ambon coastal waters (Table
Station | Collection dates | Coordinates |
---|---|---|
Alang | 14 March 2009, 19 March 2014 | 3°46'18.2"S, 128°00'24.6"E |
Lilibooi | 19 March 2014 | 3°45'08.8"S, 128°01'24.6"E |
Suli | 18–19 March 2014 | 3°37'38.2"S, 128°18'25.0"E |
Hutumuri | 14 March 2009, 19 March 2014 | 3°41'27.5"S, 128°17'56.3"E |
Mahia | 18–19 March 2014 | 3°44'42.6"S, 128°11'24.6"E |
Airlouw | 14 March 2009, 19 March 2014 | 3°46'32.5"S, 128°07'53.5"E |
In the laboratory, the epitokous nereidids were separated from other species of wawo by gross differences in the body form. The primary characteristics used to identify the different types of nereidid wawo were form of the parapodia and head morphology including eyes, antennae number and proboscis form. Voucher specimens were deposited at the Museum and Art Gallery of the Northern Territory, formerly Northern Territory Museum (NTM), Darwin, Australia, the Museum Zoologicum Bogoriense (MZB), Bogor, Indonesia, and the Reference Collection LIPI Ambon (RCLA), Indonesia (RCLA belongs to the Research Center for Deep Sea, Indonesian Institute of Sciences). Comparative material was sourced from the former Zoological Museum Amsterdam (ZMA) and Naturalis (formerly Rijksmuseum van Natuurlijke Historie, RMNH), Leiden; the ZMA collection is now integrated into the RMNH collection.
Nereidid samples collected by D.S. Hoedt and V.D. Velde were studied during a visit to Naturalis (RMNH), Leiden, in 2009 by CJG. The former comprises hundreds of epitokous Nereididae in one jar (Fig.
Preserved specimens were examined using stereo (Nikon SMZ 1500 and Nikon SMZ 645) and compound (Nikon ECLIPSE 80i and Nikon ECLIPSE 50i) light microscopes. Macrophotographs of preserved animals were taken with a Canon 5D Mark II with a Canon MPE-65 Macro Lens.
Potential new species were flagged and identified using informal names or ‘cf.’; for these taxa there is currently insufficient information on morphological variation to assign a Linnean binomen. The informal species epithet takes the general format ‘colloquial name_voucher number_name of person recognising the species’, for example, Neanthes ‘sp_Ambon_NTMW19037’ Glasby. Species are arranged alphabetically by genus and species, with informally named species at the end of each genus. Terminology for parapodial features follows
Ceratonereis (Ceratonereis) singularis australis Hartmann-Schröder, 1985: 46–47, figs 48–58.
Ceratonereis (Ceratonereis) cf. singularis australis.–
Nereis (Ceratonereis) tentaculata. –
Exmouth, Western Australia, Australia.
1 female (NTM W25886), 1 male (NTM W25890), 2 females (MZB.Pol.00175), 2 females (RCLA.Ann.048), all from Suli, Ambon Island, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 8 ex. including 5 females (NTM W23808), Banda, Maluku Province, Indonesia, coll. V.D. Velde, May 1921 (donated to NTM, formerly ZMA VPol 0962).
1 ex.(NTM W22557), 1 ex.(NTM W23939), 1 ex.(NTM W23950), 1 ex.(NTM W23983), Lizard Island, northern Great Barrier Reef, Australia, coll. CREEFS surveys 2008–2010.
Female: length (11–15 mm), maximum width (2.9–3.0 mm). Male: length (25 mm), maximum width (3.1 mm).
Ceratonereis species with females having dark brown bands on chaetiger 2 and 3 dorsally, and sometimes weaker brown bands on chaetigers 5–14; pigment apparently absent in males (1 specimen examined) (Figs
The specimens examined agree well with
Ceratonereis (Ceratonereis) cf. perkinsi by
Eastern Indonesia and Australia (widespread).
Nereis (Ceratonereis) marmorata Horst, 1924: 177–178, pl. 34, figs 13–16.
between Gisser and Seram-Laut, Maluku, Indonesia.
3 males (RCLA.Ann.042), 1 female (RCLA.Ann.043), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 3 males (RCLA.Ann.044), 2 females (RCLA.Ann.045), Suli, Ambon, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 male (RCLA.Ann.046), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 19 March 2014; 3 males (MZB.Pol.00161), 2 females (MZB.Pol.00162), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 1 male (NTM W25889), 1 female (NTM W25888), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 ex. (NTM W23811), Salawati, Raja Ampat, Indonesia coll. unknown, 18 August 1899 (donated to NTM, formerly ZMA VPol 0980).
Nereis (Ceratonereis) marmorata syntypes: 17 ex. (ZMA Vpol 0869) and 5 ex.(RMNH 1352), Siboga Stn. 172, Gisser anchorage, between this island and Seram- Laut, Maluku, Indonesia, 3°53'9.2"S, 130°51'56.2"E, coll. 26 August 1899. Composetia marmorata: 1 ex.(NTM W22530), 1 ex.(NTM W22615), 1 ex.(NTM W22797), Lizard Island, northern Great Barrier Reef, Australia, coll. CREEFS surveys 2008–2010.
Male: length (20–40 mm), maximum width (3.0–5.0 mm). Female: length (17–31 mm), maximum width (2.5–4.9 mm).
Composetia species having distinctive marmorated stripes on the prostomium and anterior body (Fig.
The distinctive marmorated (=veined) stripes on this species of Composetia facilitate identification. The markings are as described by
Indonesia. Lizard Island, Australia.
Nereis unifasciata Willey, 1905: 271–272, pl. 4, figs 85–88.
Cheval Paar, Sri Lanka
4 males (RCLA.Ann.020), 6 females (RCLA.Ann.021), Alang, Ambon Island, Indonesia, 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 19 March 2014; 10 males (RCLA.Ann.022), 11 females (RCLA.Ann.023), Lilibooi, Ambon Island, Indonesia, 3°45'08.8"S, 128°01'24.6"E, coll. J. Pamungkas, 19 March 2014; 111 males (RCLA.Ann.024), 57 females (RCLA.Ann.025), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 2 females (RCLA.Ann.026), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 38 males (RCLA.Ann.027), 41 females (RCLA.Ann.028), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 18 March 2014; 6 males (RCLA.Ann.029), 5 females (RCLA.Ann.030), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 19 March 2014; 3 males (MZB.Pol.00169), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 18 March 2014; 1 male (MZB.Pol.00170), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 4 males (NTM W23791), 5 females (NTM W23792), Hutumuri, Ambon Island, Indonesia, 3°42.1'S, 128°17.5'E, coll. J. Pamungkas, 14 March 2009; 2 males (NTM W23796), Alang, Ambon Island, Indonesia, 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 14 March 2009; 1 ex. (specimens extracted from a jar of many hundred worms in the collection of Naturalis, Leiden) (NTM W23806), Ambon Island, Indonesia, coll. D.S. Hoedt, 1866.
Male: length (15–25 mm), maximum width (1.5–2.5 mm). Female: length (18–28 mm), maximum width (2.0–3.0 mm).
Neanthes species having dark brown band on dorsal surface of chaetiger 2 and lighter bands on following pre-natatory chaetigers (Fig.
Nereidid epitokes, preserved specimens, dorsal view. A Neanthes unifasciata, male B Neanthes unifasciata, female C Neanthes sp. cf. N. gisserana male, inset showing close up of pygidial rosette D Neanthes sp. cf. N. gisserana female. mdc = modified dorsal cirri. Scales bars: 2 mm (A, B), 3 mm (C), 4 mm (D).
The specimens examined in this study agree well the description of
Indo-west Pacific (widespread).
Ceratonereis pachychaeta Fauvel, 1918: 506–508, fig. 3a–h
Neanthes pachychaeta. –
Djibouti and Madagascar.
Ambon Island, 1 specimen (RMNH inreg.), collected D.S. Hoedt, 1866 [specimen extracted from a jar of many hundred worms in the collection of Naturalis, Leiden; Fig.
Not available.
This species was recently re-described by
Indo-Pacific, widespread.
Nereis (Lycoris) gisserana Horst, 1924: 151–152, pl. 30, figs 6, 7.
5 males (RCLA.Ann.031), 3 females (RCLA. Ann.032), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 1 male (RCLA.Ann.033), 1 female (RCLA.Ann.034), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 3 males (RCLA.Ann.035), 2 females (RCLA.Ann.036), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 18 March 2014; 2 males (RCLA.Ann.037), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 19 March 2014; 2 males (MZB.Pol.00167), 2 females (MZB.Pol.00168), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 1 male (NTM W23797), Alang, Ambon Island, Indonesia 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 14 March 2009; 1 female (specimen extracted from a jar of many hundred worms in the collection of Naturalis, Leiden); Fig.
Neanthes gisserana (Horst, 1924). Syntypes 2 ex.(ZMA VPol 0854), Siboga Stn. 172, Gisser anchorage, between this island and Seram-Laut, Maluku, Indonesia, 18 m, coll. 26 August 1899. Neanthes cf gisserana. 1 ex.(NTM W22501), Lizard Island, northern Great Barrier Reef, Australia, coll. CREEFS surveys 2008–2010.
Male: length (16–25 mm), maximum width (2.0–4.0 mm). Female: length (16–32 mm), maximum width (3.0–4.0 mm).
Neanthes species having brown band on dorsal surface of chaetiger 4 (Fig.
The specimens examined in this study were compared with Horst's, syntypes, which they resemble closely, especially in the paragnath count and pattern; however, the present specimens are much larger than those of Horst’s, the modified chaetigers start later in the present specimens, and Horst does not mention the presence of a brown band which we observed on chaetiger 4. Therefore our material possibly represents a new species; it is very similar to specimens also referred to as Neanthes sp. cf N. gisserana by
Nereis (Lycoris) masalacencis Grube, 1878: 75–76, pl. 5, fig. 4.
22 males (RCLA.Ann.038), 16 females (RCLA.Ann.039), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 5 males (RCLA.Ann.40), 1 female (RCLA.Ann.41), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 male (MZB.Pol.00171), 5 females (MZB.Pol.00172), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 3 males (NTM W25892), 3 females (NTM W25891), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; Ambon Island, 4 females, 2 males (NTM W23805), collected D.S. Hoedt, 1866 (specimen extracted from a jar of many hundred worms in the collection of Naturalis, Leiden; Fig.
Male: length (30–47 mm), width (2.5–3.0 mm). Female: length (30–50 mm), width (2.0–3.0 mm).
Neanthes species having uniform brown pigmentation on dorsal surface, apparently darker in male compared to female (Fig.
The present material bears a general resemblance to Grube's species, which was described from Masalac, Philippines, in terms of pigmentation pattern and paragnath arrangement and counts. Our specimens differed in lacking paragnaths in Area I (Grube illustrates 2 paragnaths) and in the relatively longer dorsal cirri (up to 3 times length of parapodia in anterior chaetigers), whereas in Grube's specimens the dorsal cirri appear to be slightly longer than the parapodia, but the angle of the illustration makes it difficult to ascertain how much longer. It probably represents a new species.
1 female (RCLA.Ann.052), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 19 March 2014; 1 male (MZB.Pol.00173), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 1 female (MZB.Pol.00174), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 male (NTM W19155), Waimahu Beach, Ambon Island, coll. Mr Talakua, 27 March 1997; 1 male (NTM W25893), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014.
Female: length (15 mm), maximum width (2.0 mm). Male: length (16 mm), maximum width (2.0 mm).
Nereis species lacking pigmentation (Fig.
This appears to be a new species, and will be described when further specimens, including atokous individuals become available. It was listed by
Nereis (Perinereis) helleri Grube, 1878: 81–82
Perinereis cultrifera. –
Perinereis helleri. –
Bohol, Philippines.
4 males (RCLA.Ann.001), 2 females (RCLA.Ann.002), Alang, Ambon Island, Indonesia, 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 19 March 2014; 6 males (RCLA.Ann.003), 6 females (RCLA.Ann.004), Hutumuri, Ambon Island, Indonesia, 3°41'27.5"S, 128°17'56.3"E, coll. E. Moniharapon, 19 March 2014; 1 female (RCLA.Ann.005), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 18 March 2014; 11 males (RCLA.Ann.006), 10 females (RCLA.Ann.007), Airlouw, Ambon Island, Indonesia, 3°46'32.5"S, 128°07'53.5"E, coll. F. E. de Soysa, 19 March 2014; 5 males (MZB.Pol.00163), 5 females (MZB.Pol.00164), Airlouw, Ambon Island, Indonesia, 3°46'32.5"S, 128°07'53.5"E, coll. F. E. de Soysa, 19 March 2014; 2 males (NTM W23794), 2 females (NTM W23795), Alang, Ambon Island, Indonesia 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 14 March 2009; 2 males (NTM W23799), 1 female (NTM W23800), Airlouw, Ambon Island, Indonesia, 3°46'32.5"S, 128°07'53.5"E, coll. J. Pamungkas, 14 March 2009; 5 ex. (specimens extracted from a jar of many hundred worms in the collection of Naturalis, Leiden); Fig.
Male: length (25–47 mm), maximum width (3.0–6.0 mm). Female: length (28–70 mm), maximum width (4.0–6.0 mm).
Large Perinereis species having brown streaky pigmentation on dorsal surface (Fig.
Nereidid epitokes, preserved specimens, dorsal view. A Perinereis helleri, male, inset shows close up of modified parapodia of natatory region B Perinereis helleri, female C Perinereis nigropunctata, male D Perinereis nigropunctata, female. mdc = modified dorsal cirri; sdc = scalloped dorsal cirri; pr = pygidial rosette. Scales bars: 2 mm (A, B), 3 mm (C, D).
The specimens examined in this study agree well the re-description of
Indo-Pacific (widespread).
Nereis nigropunctata Horst, 1889: 171, pl. 8, figs 1–3.
Perinereis cultrifera. –
Perinereis nigropunctata. –
Malaysia.
7 males (RCLA.Ann.008), 9 females (RCLA.Ann.009), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 18 March 2014; 19 males (RCLA.Ann.010), 17 females (RCLA.Ann.011), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 18 males (RCLA.Ann.012), 32 females (RCLA.Ann.013), Hutumuri, Ambon Island, Indonesia, 3°41'27.5"S, 128°17'56.3"E, coll. E. Moniharapon, 19 March 2014; 1 male (RCLA.Ann.014), 4 females (RCLA.Ann.015), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 18 March 2014; 1 male (RCLA.Ann.016), 1 female (RCLA.Ann.017), Mahia, Ambon Island, Indonesia, 3°44'42.6"S, 128°11'24.6"E, coll. A.S. Leatemia, 19 March 2014; 1 male (RCLA.Ann.018), 4 females (RCLA.Ann.019), Alang, Ambon Island, Indonesia, 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 19 March 2014; 5 males (MZB.Pol.00165), 3 females (MZB.Pol.00166), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 ex.(NTM W23793), 1 female (NTM W23798), Alang, Ambon Island, Indonesia, 3°46'18.2"S, 128°00'24.6"E, coll. J. Pamungkas, 14 March 2009; 3 females (NTM W23801), Airlouw, Ambon Island, Indonesia, 3°46'32.5"S, 128°07'53.5 E”, coll. F. E. de Soysa, 14 March 2009; 5 ex. be (specimen extracted from a jar of many hundred worms in the collection of Naturalis, Leiden) (RMNH unreg.), Ambon Island, Indonesia, coll. D.S. Hoedt, 1866.
Male: length (10–20 mm), maximum width (2.0–3.0 mm). Female: length (11–23 mm), maximum width (2.0–4.0 mm).
Perinereis species having brown, streaky pigmentation on dorsal surface (Fig.
The specimens examined in this study agree well the description of
Indo-west Pacific (widespread).
Solomononereis merauensis Gibbs, 1971: 152–153, fig. 8a–h. –
Solomon Islands.
1 female (MZB.Pol.00176), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014; 1 female (NTM W25887), Suli, Ambon Island, Indonesia, 3°37'38.2"S, 128°18'25.0"E, coll. R. Alik, 19 March 2014.
Male: not available. Female: length (20 mm), maximum width (3.0 mm).
Solomononereis species lacking body pigmentation (Fig.
The present specimen fits the type description closely.
Indonesia, Northern Australia, Solomon Islands.
Leptonereis foli Fauvel, 1930: 520, fig. 3.
Websterinereis foli. –
Nicon sp. –
Ile de Pins, New Caledonia.
The specimens reported as Nicon sp. by
Western Pacific, Indonesia, Lizard Island (Australia).
The following key provides a means of identifying male and females of each species of Nereididae known to swarm at Ambon Island. It includes species reported both in the present study and in
1 | Body with 2 regions (largely unmodified pre-natatory region and natatory region with modified parapodia and chaetae) | 5 |
– | Body with 3 regions (pre-natatory and one or two distinct natatory regions) | 2 |
2 | Single natatory region, restricted to mid-body | 3 |
– | Two natatory regions, extending to pygidium (mid-body region with modified parapodia and oval-shaped posterior region with extremely reduced parapodia) | Solomononereis merauensis |
3 | Pre-natatory region with up to 36 chaetigers | 4 |
– | Pre-natatory region with 37–40 chaetigers; unpigmented (females) or head and anterior segments red-brown lines and spots (males) | Websterinereis foli |
4 | Pre-natatory region with 14 chaetigers (female); brown pigment restricted to head region | Ceratonereis sp. cf. C. perkinsi |
– | Pre-natatory region with 16–17 chaetigers (male and female); females with dark brown bands on chaetigers 2 and 3 | Ceratonereis singularis australis |
– | Pre-natatory region with 28–33 (male) or 34–36 (female) chaetigers; brown pigment restricted to head and pygidium | Ceratonereis sp. sensu |
5 | Dark brown band on dorsal surface of chaetiger 2 (fainter brown bands on subsequent chaetigers); anterior region with 16 (male) and 19–20 (female) unmodified chaetigers | Neanthes unifasciata |
– | Dark brown band on dorsal surface of chaetiger 4; anterior region with 21–22 (male) and 26 (female) unmodified chaetigers | Neanthes sp. cf. N. gisserana |
– | Body pigmentation not as above, or absent | 6 |
6 | Neuropodia of natatory region with distinctive ramified lamellae | Neanthes pachychaeta |
– | Neuropodia of natatory region with smooth-edged lamellae | 7 |
7 | Female epitokes with 17 or 18 pre-natatory region chaetigers | 8 |
– | Female epitokes with 19 or 20 pre-natatory region chaetigers (male 17) | Perinereis helleri |
– | Female epitokes with about 26 pre-natatory region chaetigers (male 22–23) | Nereis sp_Ambon_NTMW19037 |
– | Female epitokes with about 32 pre-natatory region chaetigers (male 28–30) | Neanthes cf. masalacensis |
8 | Male epitokes with about 14 pre-natatory region chaetigers (female 17) | Perinereis nigropunctata |
– | Male epitokes with 17 or 18 pre-natatory region chaetigers (female also 17–18) | Composetia marmorata |
Taxonomic utility of epitokal modifications. Sexual dimorphism among reproductive Nereididae is well known – it includes differences in the number of basally-swollen anterior dorsal and ventral cirri, the number of pre-natatory region chaetigers, and the presence of undersurface scalloping on the natatory dorsal cirri and pygidial rosettes in males only (
Less well known is that Nereididae appears to exhibit epitokal modification patterns at the generic level. Combining data from this study with literature information on Platynereis (
Finally, some caution must be attributed to the generality of these patterns of epitokous modifications as possibly they are dependent on the state of sexual maturity of the individual and, of course, they are based on observations of very few species per genus.
How many species? The number of species recognised as comprising wawo in Maluku is increasing as we collect more intensively and overcome taxonomic problems. In the first publication on wawo,
Rumphius (1999) describes one form that resembles a millipede:
… about the thickness of an oaten pipe, quite like young Millipedes, of a mixed green, brown, and white, and look indeed somewhat disgusting, but these have a special name, and are not considered the true Wawo.
And further, that the millipede form:
… illumes at night, giving off a clear light, which makes people avoid them even more, since they share this attribute with the Millipedes.
The millipede form may represent the shorter-bodied nereidids with their prominent parapodia; however, luminescence has, as far as we know, not been reported previously in swarming members of this family, so doubt must remain over the identity of Rumphius's, ‘millipedes’. The more abundant true wawo referred to by Rumphius (1999) as ‘like silken Floss, all entangled in small clumps’ are most likely Palola viridis and possibly other eunicid species (see
Comparison of swarming nereidid species collected by Hoedt (1866), reported by
Species | 1866 | 1995 | 2009/2014 |
---|---|---|---|
Ceratonereis sp. cf. C. perkinsi | Absent | Present | Absent |
Ceratonereis singularis australis | Present | Present | Present |
Ceratonereis sp_Martens et al. | No | Present | Absent |
Composetia marmorata | Absent | Absent | Present |
Neanthes sp. cf. N. gisserana | Present | Absent | Present |
Neanthes sp. cf. N. masalacensis | Absent | Absent | Present |
Neanthes pachychaeta | Present | Absent | Absent |
Neanthes unifasciata | Present | Present | Present |
Nereis sp_Ambon_NTMW19037 | Absent | Absent | Present |
Perinereis helleri | Present | Present | Present |
Perinereis nigropunctata | Present | Present | Present |
Solomononereis merauensis | Absent | Absent | Present |
Websterinereis foli (=Nicon sp. sensu |
Absent | Present | Absent |
Finally, it is worth noting that wawo have been reported to swarm in islands nearby Ambon, such as Haruku and Nusalaut (JP pers. obs.) and therefore are probably widespread in Maluku Province. In the Banda Islands (south-east of Ambon Island), the natives call them ‘oele’. Further, studies of Nereididae by
We are grateful to Hanung A. Mulyadi, Daniel D. Pelasula and Wahyu Purbiantoro for facilitating this study. We would also like to thank our colleagues who helped us a lot in collecting wawo specimens, i.e. Abraham S. Leatemia, Eduard Moniharapon, Franky E. de Soysa, Sem Likumahua and Robert Alik. CG would like to thank Bert Hoeksema (Naturalis, Leiden) for facilitating his study of Nereididae at Naturalis in 2009 under a Temminck Fellowship, which has led to a far greater understanding of the family in the Indo-west Pacific region. Finally, we thank Robin Sanchez-Alt and Alyson Malpartida for editing images and preparing the taxonomic plates and two anonymous referees for suggestions resulting in significant improvements.