ZOOLOGIA 31 (3): 264–270, June, 2014
http://dx.doi.org/10.1590/S1984-46702014000300008
Reinstatement of Euarche rudipalpa (Polychaeta: Acoetidae), with
remarks on morphology and body pigmentation
Sergio I. Salazar-Vallejo1,2,4, Alexandra E. Rizzo2 & Marcelo V. Fukuda3
1
Estructura y Función del Bentos, Departamento de Sistemática y Ecología Acuática, ECOSUR. Chetumal, Quintana Roo,
México. E-mail: ssalazar@ecosur.mx
2
Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro. Maracanã, 20550-900 Rio de
Janeiro, RJ, Brazil E-mail: aerizzo@hotmail.com
3
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo. 05508-090 São Paulo, SP, Brazil.
E-mail: fukuda@ib.usp.br
4
Corresponding author.
ABSTRACT. Eupanthalis rudipalpa Amaral & Nonato, 1984 was described based on three incomplete, colorless specimens and has not been reported since its original description; the epithet rudipalpa means “hirsute palps”, a characteristic that separates it from other congeneric species. Eupanthalis McIntosh, 1876 and Euarche Ehlers, 1887, and their
respective type species, E. kinbergi McIntosh, 1876 and E. tubifex Ehlers, 1887, were considered synonyms (E. rudipalpa
as a junior synonym of E. tubifex, a cosmopolitan species) by some authors, because both have sessile eyes and two or
three antennae. Nonetheless, these two genera are, in fact, distinct. Their type species can clearly be separated using
some prostomial features. We provide an emended diagnosis of Euarche, and re-instate E. rudipalpa, redescribing it
based on a Brazilian specimen (original type specimens lost). Illustrations showing body pigmentation pattern and a
key to all species of Eupanthalis are also provided.
KEY WORDS. Acoetids; color pattern; Eupanthalis; key to species; Polychaetes.
Acoetid polychaetes are large, tubicolous worms that
usually appear in very low densities in oceans; in most cases,
species have been described based on single specimens and have
seldom been recorded again (BUCHANAN 1894, DARBOUX 1899,
FAUVEL 1914a, PETTIBONE 1989). AMARAL & NONATO (1984) described
Eupanthalis rudipalpa based on three incomplete, colorless specimens, and named it because it has hirsute palps, which separate it from other species. Probably following FAUCHALD (1977),
they assigned their new species to Eupanthalis McIntosh, 1876.
FAUCHALD (1977) was following the revision of the family by
BUCHANAN (1894), who regarded Euarche Ehlers, 1887 as a junior synonym of Eupanthalis.
Eupanthalis was proposed by MCINTOSH (1876), with E.
kinbergi as its type species. It is characterized by sessile eyes,
whereas the other known genera have pedunculate eyes (BAIRD
1868, GRUBE 1876), and by having a pair of lateral antennae,
but no median one: “There is no tentacle (= median antenna)
in the specimen; but on each side of the median groove there
is a filiform or slightly subulate antenna.” (MCINTOSH 1876).
Euarche was proposed by EHLERS (1887), with Euarche tubifex as
the type species, for species that have sessile eyes (oculis non
pedunculatis) and three antennae (tentaculis 3).
BUCHANAN (1894) reviewed Acoetidae and diagnosed
Eupanthalis as having: “three prostomial tentacles, except (?)
in E. tubifex” and explained this in a footnote: “McIntosh’s
remark that there is ‘no’ median ‘tentacle in the specimen’
(actually E. kinbergi but the quote was given for E. tubifex) seems
rather to imply that there may once have been one, which has
been lost by accident.” Consequently, BUCHANAN mistook the
type species of Eupanthalis and Euarche, and assumed that McIntosh had seen a species that came to be described 11 years later.
Since these two genera were regarded as synonymous, thinking that their type species were also synonyms was probably
an unavoidable consequence, and E. tubifex was regarded as a
junior synonym of E. kinbergi by several taxonomists (FAUVEL
1914a, b, AUGENER 1918, MONRO 1928, HARTMAN 1938).
PRUVOT & RACOVITZA (1895) objected the generic arrangement proposed by BUCHANAN, but their observations were overlooked by later authors. Despite the fact that the characters
of the anterior end proposed by them have been later extensively used for separating similar genera, and their relevance
was highlighted by FAUVEL (1897): “Les meilleures caractères
pour la classification des Acoetidés restent donc ceux tirés du
nombre, de la forme et de l’implantation des appendices
céphaliques et ceux tirés des soies et des parapodes”, the best
characters for acoetid classification are in the number, shape
and insertion of cephalic appendages, and those of chaetae
and parapodia.
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Reinstatement of Euarche rudipalpa, with remarks on morphology and body pigmentation
PETTIBONE (1989) regarded Eupanthalis and Euarche as distinct and re-established Euarche, indicating that Eupanthalis has
two antennae, whereas Euarche has three. She also proposed a
further difference between the two, which is the presence and
extension of papillae along the palp’s surface, being smooth
in Eupanthalis or with papillae restricted to half their length,
whereas in Euarche palps have papillae over at least two-thirds
of their length. BARNICH & FIEGE (2003) clarified the differences
by indicating that Euarche has median antenna, and anterior
chaetigers biramous, whereas in Eupanthalis there is no median antenna, and anterior chaetigers are subbiramous, usually without notochaetae.
PETTIBONE (1989) regarded some previously described species as junior synonyms of E. tubifex, including E. rudipalpa.
Due to this expanded definition, E. tubifex ended up having a
cosmopolitan distribution. The name encompassed specimens
from the Gulf of Mexico (type locality), Gulf of Panama, Western Africa, South America, the Indian Ocean, and also specimens from Japan (IMAJIMA 1997) and the Mediterranean Sea
(BEN-ELIAHU & FIEGE 1994, BARNICH & FIEGE 2003). However, we
believe that E. tubifex is composed of different species that can
be separated by using features of the anterior end, as demonstrated in the present work. In this contribution we propose
an emended diagnosis of Euarche, a redescription of E. rudipalpa
re-establishing the taxon as a valid species; and provide illustrations based on recently collected material, showing the pigmentation pattern of the body. A key to all species of Euarche
is also provided.
MATERIAL AND METHODS
One specimen was collected in the Santos Basin, in the
Brazilian continental shelf. Some parapodia were removed and
mounted in ethanol-glycerol for semi-permanent preparations.
In one parapodium, chaetae were trimmed to observe their
arrangement in frontal view, as suggested by IMAJIMA (1997).
Some other parapodia were cross-sectioned to observe enteric
diverticula. A series of digital photographs were stacked by using
the Helicon Focus software. It is a program for focus stacking
or, in other words, a post-processing technique that extends
the depth of field in photographs. The material is deposited at
the Polychaeta Collection, from Universidade do Estado do
Rio de Janeiro (UERJ).
TAXONOMY
Euarche Ehlers, 1887
Type species. Euarche tubifex Ehlers, 1887, by monotypy.
Diagnosis (modified from BARNICH & FIEGE 2003). Prostomium with sessile eyes, rarely missing; two lateral and one
median antennae. Palps long, papillate; tentacular segment
distinct dorsally, sometimes with chaetae. Segment 2 with elytra
and long ventral cirrus. Parapodia biramous, without branchiae;
265
notopodium reduced, notochaetae delicate, long capillaries;
neuropodium projecting, neurochaetae of different kinds: long,
slender, with many long spines in distal part; short, slender
with few widespread spines; acicular with hirsute tip; lanceolate,
twisted, with many whorls of spines. Pharynx with paired
marginal papillae. Jaws 2 pairs, each with 3-8 teeth.
Remarks. Most diagnostic features are rather easy to discern; however, some notopodial features might be difficult to
evaluate, because they are often reduced to a small notoacicular
lobe, being better developed on anterior chaetigers, with
notochaetae delicate and scarce, usually emerging below the
lobe. On the middle and more posterior parapodia, notopodia
are barely developed and notochaetae are often lost. Even in
parapodial mounts, notopodia can be difficult to discern because notoacicula migrate ventrally, being placed slightly above
neurochaetal bundle bases. However, notochaetae have a
slightly different orientation, being directed obliquely from
the bundle, whereas neurochaetae tend to have a more or less
parallel orientation.
Euarche rudipalpa (Amaral & Nonato, 1984)
Figs 1-17
Eupanthalis rudipalpa Amaral & Nonato, 1984: 14, figs 1-11;
Amaral et al., 2013: 18.
Euarche tubifex: Pettibone, 1989: 14 [part].
Material examined. One specimen (UERJ 4024), Santos
Basin (28°09’57.6"S, 48°33’50.4”W to 24°51’39.6"S, 40°36’39.6”W)
Station 17, 2013 m.
Body subcylindrical anteriorly, depressed medially and
posteriorly, with wide, brownish dorsal band (Fig. 1), extending up to chaetiger 30, gradually narrowing, becoming finer
and reaching posterior end (Fig. 5). Parapodia whitish, dorsally with two or three thin, transverse pigmented bands, ventrally whitish. Venter with deep longitudinal furrow reaching
posterior end, with fine tiny brownish spots throughout; ventral cirrophores with pigmented glands (Fig. 6). Everted pharynx maculated, lateral papillae darker distally; marginal papillae
pale, several with darker inner glands; jaws dark brown (Fig.
4). Palps pale brown, blunt, darker sub-distally, tips whitish,
with longitudinal series of dark papillae running throughout
their length, and several irregular, thin, brown rings (Figs 2,
3). Antennae and tentacular cirri maculated, darker or with
larger spots ventrally. Prostomium and surrounding areas
brownish, with shiny bluish hue dorsally, along a few anterior
chaetigers (Fig. 3). Tentacular cirrophores dark brown, with
digitiform projection near cirrostyle, cirrostyles paler, with
some dark brown spots irregularly distributed. Prostomium
oval, wider than long. Anterior eyes larger, half-moon shaped,
posterior ones smaller, regularly distributed over prostomial
surface. Lateral antennae on anterior prostomial margin, tapering, tips thin, pointed, brownish becoming paler distally;
median antenna about as long as, or slightly longer than, lateral antennae, with ceratophore distinct, basal region darker
ZOOLOGIA 31 (3): 264–270, June, 2014
S.I. Salazar-Vallejo et al.
266
1
3
2
4
5
6
Figures 1-6. Euarche rudipalpa, (UERJ 4024): (1) anterior fragment in lateral view, posterior fragment in dorsal view; (2) anterior end,
dorsal view; (3) same, without elytra from chaetiger 1 (chaetae of tentacular segment too transparent and delicate to be captured); (4)
anterior end, lateral view; (5) posterior end, dorsal view; (6) same, ventral view. Scale bars: 1= 11.3 mm, 2 = 3.3 mm, 3 = 0.9 mm, 4 =
3.1 mm, 5 = 0.4 m, 6 = 0.3 mm.
than distal half (whitish). Palps about three times longer than
lateral antennae, about one-fourth longer than tentacular cirri.
Tentacular segment (first chaetiger) reduced with abundant,
delicate notochaetae and a notoaciculum. Second chaetiger
with biramous parapodia and first pair of elytra. All parapodia
biramous; anterior parapodia (Fig. 7) more prominent and pigmented than median (Fig. 8), or posterior ones (Fig. 9). Dorsal
cirri globose basally,tapering distally to a fine tip, longer than
neurochaetal fascicles in first 2-3 chaetigers, smaller in following chaetigers, barely reaching neurochaetal lobes in median
or posterior chaetigers. Notochaetae reduced to few, delicate,
smooth capillaries, decreasing in abundance and size in median and posterior chaetigers, situated ver y close to
neurochaetal fascicles. Neuropodia with larger postchaetal lobe,
margin more heavily pigmented; ventral cirri medially swollen, tapering to delicate tips, longer in first 3-4 chaetigers,
smaller medially and posteriorly. Neurochaetae arranged in
distinct series: larger neurochaetae in more or less radial series;
smaller, lower chaetae in several series in a U-shaped pattern
(Figs 4, 10). Superior supracicular neurochaetae arranged in a
ZOOLOGIA 31 (3): 264–270, June, 2014
single series, of two kinds: long, slender with many long spines
in distal part (Fig. 11) and shorter, with widespread spines;
middle neurochaetae arranged in 5-7 series, acicular, penicillate, tip falcate, bidentate, with additional fibers giving an hirsute appearance (in middle and posterior parapodia 2-3 chaetae
per series) (Fig. 12, insert); inferior neurochaetae in 3-4 series,
lanceolate, twisted and medially swollen, with numerous
whorls of spines (Fig. 13). Pharynx with 11 pairs of marginal
papillae (Fig. 14), mid-ventral and mid-dorsal ones slightly
longer; mid-dorsal papilla entire, mid-ventral papilla trilobed.
On pharynx sides, two depressed, semicircular large basal papillae, and in the inner cavity, three accessory inner papillae
dorsally and ventrally, some with a blunt projection. Jaws not
fully exposed; teeth not visible. Insertion of elytra is on parapodia 2, 4, 5, 7,... until the end of the body. First pair of elytra
large, ovoid, covering prostomium; following decreasing rapidly in size, small oval by chaetiger 10, continuing to end of
body. Elytra delicate, margin smooth (Fig. 16), surface with
rounded microtubercles, more conspicuously towards posterior body (Fig. 17), and variable amounts of brownish spots, in
Reinstatement of Euarche rudipalpa, with remarks on morphology and body pigmentation
7
10
8
11
267
9
12
13
Figures 7-13. Euarche rudipalpa, (UERJ 4024): (7) chaetiger 17, right parapodium, posterior view; (8) chaetiger 42, right parapodium,
anterior view; (9) chaetiger 98, right parapodium, anterior view with a dark orange oocyte; (10) chaetiger 60, left parapodium, frontal
view, hirsute chaetae trimmed showing chaetal serial arrangement; (11) Chaetiger 17, upper neurochaetae with uniseriate chaetae;
(12) chaetiger 42, hirsute chaetae (insert: close-up of tip); (13) same, twisted spinulose chaetae. Scale bars: 7 = 0.6 mm, 8 = 0.8 mm,
9 = 0.6 mm, 10 = 170 µm, 11 = 50 µm, 12 = 65 µm, 13 = 70 µm.
some elytra especially abundant towards elytral margin. Pygidium damaged; anal cirri unknown.
Internal features: Enteron with lateral diverticula, penetrating parapodia as large caeca, visible by transparency as
blunt opaque lobes (Fig. 9), with thick walls (Fig. 15), starting
from about chaetiger 15. Dark-orange to brownish oocytes
sparse in some posterior chaetigers, pale in ovary, dark orange
or brownish once detached, reaching about 250 µm in diameter (Fig. 9).
Distribution. Brazil, off state of Rio Grande do Sul and
Santos Basin, at 195 m depth in medium sand (in original description), now extended to deep water (2013 m).
Remarks. This specimen mostly fits the original description of Eupanthalis rudipalpa Amaral & Nonato, 1984 but di-
rect comparison with type material was not possible because
the type specimens seem to be lost (Mônica Petti, pers. com.).
Due to the presence of three antennae, and palps with papillae throughout their length, the species must be transferred to
Euarche Ehlers, 1887, as already indicated by PETTIBONE (1989:
14). Eupanthalis rudipalpa must be reinstated because it clearly
differs from other species in the genus, as shown in the key
below. Euarche rudipalpa resembles E. tubifex in the size of the
anterior eyes, which are about twice as large as the posterior
ones. This similarity might explain why the two species were
regarded as synonyms. However, E. rudipalpa and E. tubifex differ in the relative size of antennae and palps, and in the relative extent of papillae series along palps. In E. rudipalpa, papillae
extend throughout the palps, and palps are about three times
ZOOLOGIA 31 (3): 264–270, June, 2014
S.I. Salazar-Vallejo et al.
268
elytron
IP
ED
MvP
ED
BP
14
MVD
VC
15
*
16
17
Figures 14-17. Euarche rudipalpa, (UERJ 4024): (14) pharynx, oblique ventral view; (15) chaetiger 60, posterior view, transverse cut; (16)
right elytron from chaetiger 8, frontal view (*enlarged area in Fig. 17); (17) same, close-up of dorsal, posterior, marginal area showing
brownish spots and microtubercles. (BP) basal papilla, (ED) enteric diverticulum, (IP) internal papilla, (MVD) mid-ventral depression,
(MvP) mid-ventral papilla, (VC) ventral cirrus. Scale bars: 14 = 1.7 mm, 15 = 1 mm, 16 = 0.5 mm, 17 = 65 µm.
longer than lateral antennae, whereas in E. tubifex papillae
extend on the distal two-thirds of palps, and palps are about
twice as long as lateral antennae.
Recent compilations (HUTCHINGS 2000) have mentioned
the presence of enteric diverticula in the parapodia of Acoetidae;
however, only one plate containing some illustrations is available for this feature. It was provided by DELLE CHIAJE (1841b: pl.
99) for Polyodontes maxillosa (Ranzani, 1817), without any explanation for the figures. The author only provided a Latin
diagnosis for the species. These illustrations were not reproduced from earlier works, including the original description
(RANZANI 1817), and show complex enteric diverticula penetrating the parapodia, separated by constricted tunnels. Each diverticulum has lateral and distal sacs per segment; two other
globose structures were regarded as testis, perhaps, because they
were indicated with a letter “t”. The second illustration was
provided by STORCH (1968: 268, fig. 4) for a cross section of an
anterior segment of Panthalis oerstedi Kinberg, 1856, but he
provided little information about lateral caeca. In E. rudipalpa,
ZOOLOGIA 31 (3): 264–270, June, 2014
lateral connections were not observed probably because the
gut was too contracted, but their extensions inside parapodia
are confirmed. MCINTOSH (1900: 402) indicated that in P. oerstedi
enteric caeca were “large and elliptical, with a narrow neck”,
and this was illustrated by STORCH (1968).
Key to species of Euarche Ehlers, 18871 (modified
from PETTIBONE 1989)
1. Prostomium oval, without anterior furrow; eyes present . 2
1’. Prostomium bilobed, with anterior furrow; eyes absent; palps
4.0-4.5 longer than lateral antennae (Gulf of Mexico) .....
............................................... E. mexicana Pettibone, 1989
2. Anterior and posterior eyes about same size ..................... 3
2’. Anterior eyes twice as large as posterior ones .................. 5
3. Palps 3 times longer than lateral antennae ....................... 4
3’. Palps twice as long as lateral antennae; eyes on widest part
of prostomium (Mediterranean Sea) ..................................
................................ E. tubifex sensu BARNICH & FIEGE (2003)
Reinstatement of Euarche rudipalpa, with remarks on morphology and body pigmentation
3". Palps 4 times longer than lateral antennae; eyes regularly
distributed on prostomium (Philippine Islands) ..............
............................................. E. maculosa (Treadwell, 1931)
4. Eyes regularly distributed on prostomium (Congo) ...........
........................................................................... Euarche sp.
4’. Eyes displaced towards posterior part of prostomium (Pacific
Panama) ................................... E. perlae (Fauchald, 1977)2
5. Palps 2.0-2.5 times longer than lateral antennae (up to 5
times in original illustration), longitudinal rows of papillae
extended over 2/3 of palp length; antennae of about the
same length (Gulf of Mexico) ....... E. tubifex Ehlers, 18873
5’. Palps about three times longer than lateral antennae,
longitudinal rows of papillae extended throughout their
length; median and lateral antennae about same length
(Brazil [E. tubifex sensu PALMERO et al. (2008) is very similar])
............................... E. rudipalpa (Amaral & Nonato, 1984)
Key notes:
1
Euarche cristata NÚÑEZ in PALMERO et al. (2008) does not belong
to this genus because it lacks the median antenna. It should
be either a. transferred to Eupanthalis, being similar to E.
kinbergi but differing by having papillae all over the palps
(instead of having smooth palps); or b. proposed as the type
species of a new genus based on the posterior prostomial
crest, which is unique among acoetid genera.
2
Euarche cristata was regarded as another junior synonym of E.
tubifex but its status as a separate species needs to be maintained; further, E. tubifex sensu IMAJIMA (1997) from Japan
differs by having chaetiger 1 with granulose dorsal surface,
fusiform tentacular cirri, and a single series of inferior most
neurochaetae in median chaetigers.
3
BARNICH & FIEGE (2003) showed that PETTIBONE’s (1989) redescription of Eupanthalis rudipalpa was based on a mixture of
specimens and clarified the differences between Eupanthalis
and Euarche. The series upon which Eupanthalis sp. (WOLF
1984) was based included specimens that have poorly defined eyes and have palps of different sizes. We believe that
it either includes one specimen in regeneration or two different species. It cannot be keyed out. Further, PETTIBONE
(1989) included in Euarche two specimens from the Arabian
Sea, identified as Eupanthalis kinbergi by HARTMAN (1938).
Formerly deposited in the Allan Hancock Foundation, they
are now in the Los Angeles County Museum, but they might
belong to a different species. WEHE & FIEGE (2002) listed other
records for the Indian Ocean. Because these records were
not illustrated, they cannot be included in the key.
ACKNOWLEDGEMENTS
Biodiversity Heritage Library, CiNii, GoogleBooks, Internet
Archive, Ruth Barnich, Dieter Fiege both from SMF, Wagner
Magalhães (University of Hawaii), and Luis F. Carrera-Parra
(ECOSUR) provided important references for this research. Ruth
269
Barnich and an anonymous reviewer critically read earlier drafts
and made significant improvements to this final document. We
also thank Mônica Petti (IO/USP) for helping us with the Prof.
Edmundo F. Nonato biological collection. SISV had a visiting
professor scholarship from the Universidade do Estado do Rio
de Janeiro (UERJ).
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