On-line version ISSN 0034-7744
Rev. biol. trop vol.59 n.4 San José Dec. 2011
Giant Eunicid Polychaetes (Annelida) in shallow tropical and temperate seas
1. El Colegio de la Frontera Sur, Chetumal, México; firstname.lastname@example.org
2. Fac. Ciencias Biológicas, Universidad Autónoma Nuevo León, Monterrey
Some species of Eunice might reach giant size, often being longer than 2m, and they are known from tropical and temperate seas. Despite their large size and recent internet notoriety, there remain some taxonomic problems in large-sized eunicids, especially since original descriptions were brief and type materials are often missing. As a mean to encourage the solution of this situation, we review the historical progress in the taxonomy of the group, including some comments on generic and specific delineation, and recommend some critical steps to solve the current confusion. These ideally would include collecting in type localities, evaluate ontogenetic morphological changes, and generate some molecular analysis to complement the morphological approach. Rev. Biol. Trop. 59 (4): 1463-1474. Epub 2011 December 01.
Key words: Eunicidae, Eunice, Leodice, Indian Ocean, Red Sea, Pacific Ocean, Caribbean Sea, Mediterranean Sea, opinion article.
Algunas species de Eunice pueden alcanzar tamaño gigantesco, a menudo sobrepasan los 2m de largo, y se conocen de mares tropicales y templados. A pesar de su gran tamaño y de su reciente notoriedad en Internet, todavía hay problemas taxonómicos entre los eunícidos gigantes, especialmente dado que las descripciones originales fueron breves y a menudo se carece de materiales tipo. Para incentivar la solución del problema, revisamos el desarrollo histórico de la taxonomía del grupo y se incluyen algunos comentarios sobre la delineación de los géneros y de las especies y recomendamos algunos pasos críticos para alcanzar este fin. Idealmente, esto incluiría recolectar en las localidades tipo, evaluar cambios morfológicos durante la ontogenia y generar algunos análisis moleculares para complementar el enfoque morfológico.
Palabras clave: Eunicidae, Eunice, Leodice, Oceano Indico, Mar Rojo, Oceano Pacífico, Mar Caribe, Mar Mediterráneo.
Some species of Eunice can be over 3m long (Pruvot & Racovitza 1895, Uchida et al. 2009), making them the largest polychaete species and placing them among the longest benthic invertebrates. The genus is very rich in species, having about 300 available names, and making it the largest genus among polychaetes. Large-bodied eunicid polychaetes have got internet notoriety because of some impressive underwater photographs, usually showing them with a fully extended jaw-apparatus, and in preparation for a fast bite over any marine organism swimming nearby their galleries. Further, "Giant Eunicids" are frequently found in marine aquaria and often become a serious concern due to the negative effect on fishes and corals, and have been repeatedly reported in specialized Web sites. Thus, in 2009 a large specimen found in a British marine aquarium reached the news because it was 1.2m long and had been eating other residents of the aquarium. Some species in the eunicid genus Marphysa de Quatrefages, 1865 may become very large, but in this note we will only refer to members of Eunice.
Most of these specimens have been regarded as belonging to Eunice aphroditois (Pallas,1788), despite some marked differences in morphology and ecology among them (see below), and especially since the distribution apparently includes all tropical seas and even some temperate areas like the Mediterranean Sea and Japan. This species, however, was described from Sri Lanka, and this wide distribution needs clarification, especially since there might be more than one species involved under the same name. Our purpose in this essay is to provide some critical historical information together with some recommendations on how to deal with the current confusion about the "Giant Eunicids" of the world.
Since the classic synthesis by May (1992), it has been found that our knowledge about species is body-size dependent, with large sized groups being better known than smaller ones (Fautin et al. 2010, Griffiths et al. 2010). Thus, it might be expected that large sized polychaetes should have no taxonomic problems, but that is certainly not the case, especially when the original description was brief and there is no type material left. This is relevant because E. aphroditois is the formal type species for Eunice, being regarded as a senior synonym for E. gigantea (see below), and for some other names that have been regarded as cosmopolitan or pantropical species. In order to clarify the problem, and despite the recent revision by Fauchald (1992), a chronological and annotated synthesis will help understand how the current situation has been reached, and why some names deserve to be restricted, reinstated or introduced, especially since original descriptions were brief and type material is often missing. The solution, of course, in order to be formal, requires detailed studies on type, topotype and additional materials, and we must proceed on that soon, but it is not the scope for this contribution.
This confusion is remarkable because there were some indications that what was regarded as belonging to a single species (E. aphroditois), was containing at least two different branchial patterns and feeding modes. McIntosh (1885:282) compared some specimens collected in Sydney, Australia, and others from the Philippine Islands; the former had single-filament branchiae from chaetiger 6, reaching up to 30 filaments in following chaetigers, while the latter had pectinate branchiae from chaetiger 5, and reaching up to 18 filaments. Further, since he used to make dissections or cross sections to investigate anatomical features, he noticed that the intestine showed interesting differences as well. In the Australian specimen there were only algae, while the Philippine worm had mud, fragments of crustaceans, algae and some spicules (holothurian?). On the other hand, Treadwell (1922:136) noticed important differences in pigmentation and relative development of anterior appendages and branchiae, but these differences were overlooked or disregarded to separate the different forms."Bobbit worms" in the web
A search for "Bobbit worms" would retrieve many images of different eunicid species. However, it must be emphasized that the name was applied for those capable of keeping their body erect from the bottom, and of fully-opening their jaw apparatus. The images available of large eunicids from tropical and subtropical seas indicate several pigmentation patterns and a variety of habitats, from sandy or muddy sediments to rocks or sponges; they probably belong to different species but none has been identified to species. These organisms could be separated on the basis of color patterns and some other morphologic traits, but it is remarkable that these differences have been overlooked by previous authors. Of course, they have not had access to the photographs based on living specimens, which is certainly our privilege, and we think reconsideration about these differences is badly needed. We will try to explain what can be easily noticed in the available images.
The originally called "Bobbit worm" is frequently photographed and seems to be living in the Philippines and in Indonesia (Fig. 1A). Its body has wide, transverse dark bands, the anterior appendages are banded, the median antenna is about as long as peristomial width, peristomial cirri are fusiform, whitish, but does not have an anterior whitish chaetiger. Branchiae start by chaetiger five.
The second species has fewer pictures and some were taken in Australia (Fig. 1B). This species differs by having a reddish, maculated body, appendages with a solid, reddish coloration, the median antenna is two-three times longer than peristomial width, peristomial cirri are reddish, either motled or solid in color, and the chaetiger four is whitish, or is at least less heavily pigmented than neighbor chaetigers. The start of branchiae is unknown.
The fourth species, which might be called the Bengal eunicid, is very spectacular and has been taken in Indonesia; there are two photos available of it (Fig. 2A, B). In this species, the body is golden with purple, intersegmental lines and some fine, irregular, longitudinal lines running throughout the body, but the chaetiger four is whitish, instead of having the background color. The antennae are solid golden, the laterals are slightly longer than peristomial width and the peristomial cirri are dark, blunt. Branchiae start by chaetiger seven.
There are apparently two large Caribbean Sea species that have been frequently taken (Fig. 3). There are two body pigmentation patterns but none has a whitish anterior chaetiger. One is dark purple, with irregular, fine, longitudinal lines in all segments, but appendages are banded (Fig. 3A, B); peristomial cirri are cirriform, banded, and not reaching the anterior margin of peristomium. Branchiae are pectinate, starting from chaetiger seven.
The other Caribbean form is dark reddish with a barely noticeable banding in anterior appendages, making them look like having a solid pigmentation (Fig. 3C-E), peristomial cirri are cirriform, not reaching the peristomial anterior margin. Branchiae are pectinate, starting in chaetiger nine. The pigmentation pattern seems to change from deep purple or violet in smaller specimens to reddish in older ones, and this is shown by the regeneration of the posterior end of a large eunicid (Fig. 3E), where the color is more violet in younger or recently formed segments.
In the following paragraphs, we introduce some comments on the status of genera. They might seem of little relevance for giant eunicids, but they illustrate the problematic situation in the family. The genera Eunice Cuvier, 1817, was proposed without naming any type species (Cuvier 1817:524-525), but after a 1.2m long specimen from the Indian Ocean. Cuvier listed as a footnote what might have been regarded as the type species (Nereis aphroditois Pallas, 1788), as indicated by Fauchald (1992:4) but it was subsequently designated by Verrill (1900). On the other hand, Kinberg (1865:561) used the relative development of the anterior prostomial margin to propose a new genus and separated several species in Eriphyle, with Eriphyle capensis Kinberg, 1865 as the type species. This genus especially contains those large-sized specimens having tetra-lobed prostomia, including E. gigantea, but excluding the remaining species, which would remain in Eunice. By the way, the type species name had to be replaced to E. kinbergi Ehlers, 1868 (:306, footnote), which might be a junior synonym of E. macrobranchia Schmarda (1861) (Text figs. a-f, Fig. 258).
This proposal by Kinberg can now be regarded as incorrect after the principle of coordination of the genus group (ICZN 1999), because the type species is the only one which could not be taken out of its corresponding genus. However, Malmgren (1867:64) indicated that following Kinberg’s proposal, Eunice should be split in two genera and because the type species was already included in Eunice, he proposed to retain Leodice Savigny, 1822, and designated L. antennata Savigny (1822), as its type species. This implied that Eunice sensu Kinberg (non Cuvier) was the same as Leodice Savigny, but the latter was restricted to exclude the type species for Eunice and other similar species. The two groups based upon these type species are rather different and well-defined, and similar species would be easily assigned to each of them; however, there are some other morphological patterns which might be more difficult to ascertain, and it deserves a deeper study. Regretfully, Malmgren’s proposal was misunderstood since Verrill (1900), resulting in either both names being used indistinctly, Leodice being regarded as a junior synonym of Eunice, while some authors (Grube, Gravier) employed Eriphyle as a subgenus.
On the other hand, Verrill (1900:650) proposed Mayeria for Staurocephalus gregaricus Mayer, 1900, which had been collected swarming in Southern Florida. However, Mayer made a clarification (Mayer 1902) indicating that his original description had been based upon posterior fragments. At the same time, he indicated that the species regarded as the Atlantic palolo was Eunice fucata Ehlers, 1885, and this clarification was repeated by Treadwell (1921: 43). It is noteworthy that Augener (1925:28) has indicated that this is a junior synonym of E. schemacephala Schmarda, 1861, which has been frequently cited in the Grand Caribbean region, but Fauchald (1992:298) regarded as indeterminable.
The species. For this section, we deal with several species either allied to E. aphroditois, or that have been regarded as its junior synonyms. The sequence is chronological and some critical comments have been inserted to help track the confusion.
Savigny (in Lamarck 1818, 1835:561-562, Savigny 1822:49) introduced Leodice gigantean for a very large specimen deposited in the Paris Museum, collected in the Indian Ocean, probably from La Reunion. Savigny included the previous description by Pallas, and the brief indication by Cuvier, which might imply they were synonyms, but Savigny had a different species. He noticed branchiae start by chaetiger five, being pectinate throughout the body with up to 35 filaments, and he also noticed that they were larger than dorsal cirri. He also indicated that because of the four-lobed head, this species was distinct from the other species and perhaps deserved to be placed in a separate group (Savigny 1822:50, Observ.).
Anyway, the correct name for the type species of Eunice should be cited as Leodice gigantea Savigny in Lamarck, 1818 (ICZN 1999). The branchial features might be different enough as to keep E. aphroditois and E. gigantea Savigny in Lamarck as distinct, but the contrary perspective has been repeatedly indicated (see below).
The combination E. gigantea Cuvier (1830:199-200), introduced for a large specimen from the Caribbean Sea, does not have priority, and might be regarded as a nomen nudum. The name was made available by Milne-Edwards (1836:32, Pl. 10, Fig. 1), but it might be called either E. gigantea Cuvier in Milne-Edwards, 1836, or E. gigantea Milne-Edwards, 1836. Anyway, since this is a junior homonym, the name must be replaced (ICZN 1999), and it might be conspecific with E. violaceomaculata Ehlers, 1887 (see below).
The next name for "Giant Eunicids" was introduced for a Mediterranean Sea species, Eunice gigantea (delle Chiaje 1825:389-393, 424, Pl. 27, Figs. 1-8). His 1.5m long specimen was reddish with yellow dorsal cirri and some bluish spots or black streaks; its antennae were subequal with pectinate branchiae starting by chaetiger 11. However, since this combination had already been used before (L. gigantea Savigny in Lamarck, 1818, L. gigantean Savigny, 1822), de Quatrefages (1866:330) proposed E. maxima as a replacement name for the species described from the Gulf of Naples. This was followed by Ehlers (1868:311), who kept the species separated from the similar E. aphroditois, but Fauvel (1923:403) regarded it as a junior synonym of E. roussaei (see below), and later (Fauvel 1932) as a junior synonym of E. aphroditois. Eunice purpurea Grube, 1866 was later described from the Adriatic Sea with a smaller specimen (Fauchald 1992:279), and seems to be closely allied to E. maxima.
On the other hand, with the materials collected by Ørsted, a large eunicid species was described from the Pacific coast of Central America (E. violacea Grube & Ørsted in Grube, 1856). Monro (1933:58) regarded this species as a junior synonym of E. aphroditois, although the pigmentation pattern, given by Crossland (who had collected the specimens and seen them alive), resembles what was originally indicated for E. violacea. This pattern is rather unique by having a purple background, solid in juveniles or in regenerating posterior regions, and having dorsal cirri with sulfuryellow bands. However, the combination of a brief description and the lack of type material implied that the species was regarded as indeterminable (Fauchald 1992:335). We think that the species deserves to be reinstated by finding topotype materials.
Ehlers (1868:309) noticed that there was confusion because the specimens in the Paris museum were coming from the Caribbean Sea and from the Indian Ocean. He accepted some differences in pigmentation and in the shape of the jaws, as illustrated by de Quatrefages for his E. roussaei, which was supposedly described from the Caribbean Sea and from the Bay of Biscay. These disparate localities deserve some further comments and are given below.
One of the Caribbean forms was described by Ehlers (1887:86-87, Pl. 24, Figs. 11, 12, Pl. 25, Figs. 1-7 as E. violaceomaculata with some specimens from Dry Tortugas, Florida. The species includes 20cm long specimens being violet or maculated, brownish-violet, with banded appendages (antennae, palps, peristomial cirri). He noticed that in almost complete concordance with the previous observations by Grube for the Martinique specimens, branchiae were pectinate, and started in chaetigers six-nine, reaching up to 40 filaments. It is noteworthy that two other similar species were described with smaller specimens from nearby localities, and might have priority over Ehlers name: E. nigricans Schmarda, 1861 (p. 131, Text figs. a-f), from Jamaica, and E. longisetis Webster, 1884 (p. 317-318, Pl. 10, Figs. 46-49 from Bermuda. Treadwell (1921:27) preferred to use the latter name, but if they are synonyms, E. nigricans would have priority.
On the other hand, the first record of E. aphroditois for the Western Pacific Ocean was made by Grube (1878:144) for the Philippine Islands. He found that his specimens were maculate brownish or dark purple, had smooth antennae, palps and peristomial cirri, pectinate branchiae always from chaetiger five, with up to 38-43 filaments, often markedly smaller than dorsal cirri. For the remarks, however, he repeated previous findings by Ehlers, who indicated that there were several specimens from Sidney, Australia, or other comments made by Kinberg from unspecified localities in the Indian Ocean, who had found that branchiae could start from chaetiger eight or nine, and in these cases branchiae were starting as single filaments. Some other studies would confirm the presence of this species in the same region. McIntosh (1885:282) corroborated the start of branchiae by chaetiger five, but his specimens had fewer filaments, probably because they were smaller.
About the large species found in the French Atlantic coast, de Saint-Joseph preferred to use E. kinbergi instead of E. roussaei, despite the fact that the latter has priority but he was following von Marenzeller (1888:7); further, by comparing Mediterranean and Vietnamese specimens, he indicated (1898:261-262) that the main differences were the relative size of the compound falciger blade, the relative length of antennae, peristomial and dorsal cirri, the first chaetiger with branchiae, and the presence of the whitish segment. However, for some species described from the same region, he thought there was a progression of pectinate branchiae, but in juveniles they were supposed to start as single filaments in chaetigers seven-ten, becoming replaced by pectinate branchiae by chaetigers 11 or 12. On the basis of this approach, he regarded as junior synonyms some other species. This deserves a detailed analysis but based upon specimens from the same locality.
Hartman (1944:110) followed Fauvel (1932) by indicating a cosmopolitan distribution for E. aphroditois, confirmed the records by Monro for the Eastern Pacific, extended its distribution from Baja California to Colombia, and following Fauvel (1934:30), even attached E. violaceomaculata as a junior synonym for E. aphroditois. Later authors did not significantly modify this perspective.
Where are we now?
As it has been noticed above, there are differences in branchial features that have been overlooked or disregarded as usable specific features, that could help us solve the confusion. At the same time, we might have to reconsider the early proposal by Pruvot & Racovitza (1895:422) about the specific characters such as pigmentation pattern, length of antennae, their relative articulation, start of branchiae and their maximal filament number (size-dependent), type of pectinate chaetae, blade relative size features in compound falcigers, form and shape of the subacicular hooks, number of abranchiate posterior chaetigers, and number and length of anal cirri.
On the other hand, it seems that the first indication of ontogenetic changes in pigmentation was given by de Saint-Joseph (1898:265-266), while those related to other features was given by Monro (1924:53). Thus, de Saint-Joseph thought that the purple pigmentation, together with the banded pattern in anterior appendages and the whitish anterior segment would gradually disappear by taking the adult pigmentation, resulting in fading off the white collar as well as any banding in the anterior appendages. However, Pruvot & Racovitza (1895:407-408) noticed that their huge specimens retained the white collar, and that antennae, peristomial and anal cirri retained the banding, while it was just retained on the dorsal cirri of few anterior chaetigers. Additional studies on growth and ontogenetic changes are needed to clarify this issue; some histological studies would show a difference in glandular pattern and a C01-barcoding would clarify the genetic affinities. Further, for what Monro regarded as juveniles of E. antennata Savigny, 1822, the antennae are more markedly moniliform, the prostomium is more prominent and the peristomium does not cover the eyes. However, the most interesting conclusion refers to the branchiae; he noticed that their start was slightly variable since in 78 specimens, they appeared mostly in chaetiger five, but in nine other specimens, they started by chaetiger four, while the number of filaments varied more markedly (4-14 filaments) and depending on body size.
Miura (1986) made a careful analysis of branchial features for many Japanese species, including what he regarded as E. aphroditois, and he studied their start and number of filaments. He noticed that in those species having branchiae starting before chaetiger nine, the first chaetiger with branchiae was size-independent, usually having a very slight variation of up to a single chaetiger, if any was present at all; however, the number of filaments was sizedependent. These findings were corroborated by Fauchald (1991) in a study about Caribbean eunicids; together with some other features of the anterior end appendages, these features have been incorporated in a modified key to all species in the world (Carrera-Parra & Salazar-Vallejo 1998).
Zanol & Bettoso (2006) have recently addressed some of the problematic, large-sized species, especially since Fauvel (1932:133) regarded E. roussaei as a junior synonym of E. aphroditois. They compared Mediterranean with Indian Ocean specimens and, on these grounds, concluded that there were two different species. However, their description of the Mediterranean specimens shows morphological differences between specimens from the Gulf of Naples and those from the Adriatic Sea, especially regarding the articulation of the anterior end appendages and the relative size of branchiae and dorsal cirri: the Adriatic Sea resembles E. purpurea as redescribed by Fauchald (1992:279). Thus, a previously regarded cosmopolitan species, Eunice aphroditois (Pallas, 1788) should be regarded as restricted to the Indian Ocean and Western Pacific Ocean, but as has been shown above, there are several different color morphs or species in the region. Anyway, this restriction is a certain improvement, and it will help clarify any forthcoming studies for the large eunicids living in the Indo-Pacific region.
On the other hand, they concluded that the Mediterranean species should be called E. roussaei de Quatrefages, 1866. We question this conclusion and think that the situation deserves an additional study to clarify the situation for the Mediterranean Sea species, where several other similar species have been described, and, at the same time, to analyze the situation of the large eunicids from the Caribbean Sea, which have also been regarded as belonging to E. aphroditois.
This essay was motivated by some findings while the three of us were studying type materials in the Paris museum where Tarik Meziane kindly accepted to receive us. The monograph by Kristian Fauchald together with the support of many colleagues from museums and collections, has allowed us to understand the current confusion and to suggest some means to overcome it. José Santos Gómez, our librarian in Ecosur-Chetumal located some difficult to find publications. We wholeheartedly thank the wonderful photographers who have made their photos or comments available in the web: Aimee J. Frank, Anne Prouzet, Artour_a, Brad Brown, Budak, David Harasti, Jean-Marie Bertot, Nivram, and Sylvain le Bris. The careful review by three anonymous referees resulted in a significant improvement in this final version. Funding for this contribution was provided by Conacyt (61609), by the Universidad Autónoma de Nuevo León and El Colegio de la Frontera Sur.
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J. Angel de León-González. Fac. Ciencias Biológicas, Universidad Autónoma Nuevo León, Monterrey