Naineris aurantiaca ( Müller, 1858 )

Alvarez, Ricardo Castro, Miranda, Vinícius Da Rocha & Brasil, Ana Claudia Dos Santos, 2019, Redescription of Naineris aurantiaca (Müller, 1858) and designation of a neotype from the Brazilian coast (Annelida: Orbiniidae), Zootaxa 4571 (1), pp. 125-136 : 128-133

Naineris aurantiaca ( Müller, 1858)

Theodisca aurantiaca Müller, 1858

Material examined. Neotype: Naineris aurantiaca , ( MNRJP 1956 ), 1 spm., under rocks between oyster banks, intertidal, 27°47'2"S– 48°30'25"W, 05/October/18, Pântano do Sul Beach , Santa Catarina, Brazil.

Additional material: Naineris aurantiaca , MNRJP 1955 , 4 spms., under rocks between oyster banks, intertidal, 27°47'2"S– 48°30'25"W, 05/October/2018, Pântano do Sul Beach , Santa Catarina, Brazil ; Naineris aurantiaca , MNRJP 1957 , 38 spms (one mounted for SEM)., in algal tufts, intertidal, 23° 3'27.93"S– 43°59'28.30"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , MNRJP 1958 , 1 spm., in algal tufts, intertidal, 23° 4'59.52"S– 44° 0'27.29"W, 19/August/2016, Praia do Sino , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , MNRJP 1959 , 11 spms., in mussel bed, intertidal, 23° 4'59.52"S– 44° 0'27.29"W, 08/April/2016, Praia do Sino , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , MNRJP 1960 , 2 spms., in algal tufts, intertidal, 23° 3'27.93"S– 43°59'28.30"W, 29/July/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , MNRJP 1961 , 6 spms., in mussel bed, intertidal, 23° 3'56.68"S– 43°59'30.65"W, 12/September/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , MNRJP 1962 , 2 spms., in algal tuft, intertidal, 23° 3'56.68"S– 43°59'30.65"W, 21/ February/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21327, 7 spms., in algal tufts, intertidal, 23° 5'5.05"S– 44° 0'31.21"W, 16/April/2017, Praia do Sino , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21328, 10 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21329, 7 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21330, 10 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21331, 9 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21332, 10 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21333, 9 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil ; Naineris aurantiaca , ZUEC-Pol 21334, 1 spms., in algal tufts, intertidal, 23° 3'57.34"S– 43°59'31.35"W, 08/March/2016, Praia Grande , Ilha da Marambaia, Rio de Janeiro, Brazil .

Comparative material examined: Naineris setosa , ZUEC-Pol 3791, 5 spms., 23°37'49.03"S– 45°24'55.47"W, 08/September/2006, Cidade Beach in Caraguatatuba, São Paulo, Brazil; Naineris setosa , ZUEC-Pol 3792, 6 spms., 23°37'49.03"S– 45°24'55.47"W, 08/August/2006, Cidade Beach in Caraguatatuba, São Paulo, Brazil; Naineris sp., ZUEC-Pol 2194, 13 spms., 23°48'47.03"S– 45°24'31.20"W, 14/September/1974, Praia do Araçá, São Paulo, Brazil; Naineris cf. laevigata ZUEC-Pol 8358, 7 spms., 23°37'38.95"S– 45°23'52.22"W, 15/March/2001, rocky shore in Caraguatatuba, São Paulo, Brazil; Naineris cf. laevigata ZUEC-Pol 8309, 18 spms., 23°46'50.09"S– 45°39'52.21"W, 08/April/2001, rocky shore in Baleia, São Sebastião, São Paulo, Brazil. Naineris sp., MUZUSP 355, 5 spms., 23°47'22.55"S– 45°21'48.61"W, 19/February/1997, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil; Naineris sp. MUZUSP 376, 1 spm., 23°47'22.55"S– 45°21'48.61"W, 23/August/1990, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 567, 3 spms., 23°47'22.55"S– 45°21'48.61"W, 12/ March/1997, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 622, 1 spm., 23°45'49.04"S– 45°20'57.23"W, 25/November/1992, Praia de Barreiros, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 655, 4 spms., 23°47'22.55"S– 45°21'48.61"W, 09/December/1996, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 714, 2 spms., 23°45'49.04"S– 45°20'57.23"W, 19/September/1990, Praia de Barreiros, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 725, 3 spms., 23°47'22.55"S– 45°21'48.61"W, 25/ June/1997, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil; Naineris sp., MUZUSP 781, 1 spm., 23°45'25.57"S– 45°24'39.22"W, 20/January/1997, Praia de São Francisco, São Sebastião, São Paulo, Brazil; Naineris sp., MUZUSP 786, 1 spm., 23°48'47.24"S– 45°24'31.35"W, 27/June/1987, Praia do Araçá, São Sebastião, São Paulo, Brazil; Naineris sp., MUZUSP 803, 2 spms., 23°47'22.55"S– 45°21'48.61"W, 18/September/1990, Praia Engenho d'Água, Ilhabela, São Paulo, Brazil., Naineris grubei australis Hartman, 1957 , holotype LACM—AHF Poly 676, 1 spm., 35°9'0"S– 138°28'0"E, Port Noarlunga, Adelaide, Australia; Naineris uncinata Hartman, 1957 ., holotype LACM –AHF Poly 675, 1 spm., 43°10'26"N– 124°19'02"W, Loss bay, Oregon, USA, 02/August/1942; Naineris bicornis Hartman, 1951 , holotype LACM—AHF Poly 674, 1 spm., 29°54'36"N– 84°23'43"W, Aligator Harbour, Florida, USA, 04/February /1950.

Measurements. NEOTYPE: length 4.84 mm (83 chaetigers), width 0.3 mm.

Additional material measured: 60 spms. from Praia Grande in Marambaia Island, Rio de Janeiro): mean length 14.45 mm (SD ± 5.87 mm), ranging from 6.36 mm (60 chaetigers) to 34.74 mm (153 chaetigers); mean width: 0.75 mm (SD ± 0.27mm), ranging from 0.21 mm to 1.42 mm; number of chaetigers (range = 50 to 153).

Diagnosis. Pre- and post-branchial papillae and interramal papillae occur only on abdominal segments. Dorsal organs occur posteriorly from thoracic chaetiger 12 onwards. Thoracic neuropodia with crenulated capillary chaetae arranged in oblique ventral rows, an oblique dorsal row of blunt-tipped uncini with lateral notches or grooves, subuluncini arranged anteriorly in 2-3 transverse rows, and a posterior transverse row of heavily-ridged uncini.

Description (Based on neotype). Prostomium rounded anteriorly, eyes absent, nuchal organ present ( Figs. 3A , 4A ). Peristomium a single achaetous annulus ( Figs. 3A , 4A ). Mouth opening antero-ventrally, labial lips striated ( Fig. 3B ). Proboscis formed by finger-shaped lobes, without ciliated epithelium. Branchiae beginning on chaetiger 9; morphologically simple, conical, with ciliated epithelium ( Fig. 3G ). First branchiae short (a sixth of the longest branchiae), becoming longer in abdominal region; equal to or smaller than notopodial lobe on anteriormost segments, becoming longer towards abdominal region where equal the body width, but decreasing in length over last segments ( Figs. 2A , 3B ). Papillae at the anterior and posterior base of the branchiae occurring only on abdominal segments; posterior papillae arranged almost laterally to branchiae ( Figs. 2 A-C, 3G-H). Dorsal organs occurring from chaetiger 12 onwards ( Fig. 3G ).

Thoracic segments depressed dorso-ventrally. Thorax comprising 13 chaetigers ( Fig. 2A ). Parapodia biramous. Interramal papilla absent ( Fig. 3F ). Parapodia pre-chaetal lobe not evident; post-chaetal lobes pyriform. Notopodial lobes longer than those of neuropodium ( Fig. 3F ). Notochaetae crenulate capillaries, arranged in transverse rows and forming an open semicircle; number of chaetae ranging from seven (on anterior segments) to 17 (on posteriormost thoracic segments). Neurochaetae comprising four types: subuluncini arranged anteriorly in 2-3 transverse rows (on up to 23 chaetae per fascicle); an oblique dorsal blunt-tipped uncini with lateral notch or groove on each side of uncinial apex (on up to four chaetae) ( Fig. 4C ); posterior transverse row of up to five heavily-ridged uncini; and row of up to six crenulated capillaries arranged obliquely and ventral to other types of neurochaetae ( Figs. 2D , 3F , 4D ).

Abdominal segments cylindrical. Abdomen with 70 chaetigers. Parapodia dorsally projected and biramous. As observed in thoracic region, only post-chaetal pyriform lobes evident. Notopodial lobe twice length of neuropodial lobe ( Fig. 3E ). Interramal papillae present between notopodia and neuropodia ( Figs. 2B , 3E ). Notochaetae crenulate capillaries, arranged in two transverse rows of up to 12 chaetae. A single lower furcate notochaeta, hosting variably-sized tines interspersed with thin needles (longest needles as long as half length of longest tine); shaft with transverse rows of small barbs ( Fig. 4F ). Neurochaetae crenulate capillaries (up to 10 chaetae) mingled with spines (up to four chaetae per parapodia) ( Fig. 4E ).

Pygidium not observed.

Variation in non type material examined: The numbers of thoracic chaetigers and the position of the first pair of branchiae were not fixed among the specimens we examined. There were 13 chaetigers in the thoracic region of the neotype, but this character varied from 9 to 17 chaetigers across all specimens observed from Rio de Janeiro State (60 spms.) and from 10 to 21 chaetigers in specimens from São Paulo State (38 spms.).

Proboscis of larger specimens (>100 chaetigers) with three thick branches at the base, forming two dorsal and one ventral lobe with numerous rami whereas smaller specimens (50–100 chaetigers) only have two lobes (one emerging on right and a larger one emerging ventrolaterally on left, both less branching than larger specimens, as reported by Eisig (1914)) ( Figs. 3C , 4B ).

Pygidia were observed in specimens from Marambaia Island, bearing a dorsal and a ventral pair of cirri, ventral pair being longer than dorsal pair ( Fig. 3D ). Anus dorsal, located between dorsal pair of cirri.

Habitat. Intertidal. This species occurred on sediment and in deposited sediments among algae rhizoids and in between the shells of mussel beds ( Perna perna ) and oyster banks.

Distribution. Type locality: Santa Catarina Island (Florianópolis), as Desterro in the original description.

Additional occurrence. São Paulo: São Sebastião, Caraguatatuba, Ubatuba, Ilhabela. Rio de Janeiro: Marambaia Island, Jaguanum Island.

Remarks. Müller (1858) did not specify a detailed location from where his specimens were sampled but assumed to be intertidal. Since the information in the original description of N. aurantiaca was minimal, we had to rely on Müller’s drawings for comparative analysis with our specimens. In so doing, we noted the presence of a dendritical proboscis and the number of proboscidial branches, the four anal cirri located around the anus, the abdominal chaetae predominantly being capillaries, the pyriform shape of abdominal post-chaetal lobes, and the abdominal branchiae being very similar to specimens we sampled from the type locality. We are therefore confident that we have obtained specimens that match the species described by Müller in 1858. The island of Santa Catarina is considered to be a well preserved location with no records of drastic changes on the island foreshore.

Morphologically, the species most closely related to N. aurantiaca is N. dendritica ( Kinberg, 1866) . Both species share the small size of the first branchiae compared to those of abdominal segments, the presence of prechaetal lamellae, and chaetal morphology. Naineris dendritica differs from N. aurantiaca in lacking pre- and postbranchial interramal papillae, the absence of a nuchal organ, and the number of basal lobes of the proboscis. The number of proboscis lobes in N. dendritica was not described by Blake (1996), but Moore (1909) described four lobes in N. robustus (an accepted synonym of N. dendritica (Hartman, 1948)) , whereas N. aurantiaca has three proboscis lobes.

Naineris aurantiaca differs from all other species of the genus occurring on the Brazilian coast, from intertidal to deeper waters species. In relation to N. laevigata both are similar in having interramal papillae, but differ in their chaetal arrangements in thoracic neuropodia. Naineris laevigata has subuluncini and a posterior row of unstriated uncini with or without hoods, whereas N. aurantiaca uncini lack hoods and has two easily differentiated types of uncini.

Naineris setosa ( Verrill, 1900) , also recorded from Brazil, differs from N. aurantiaca in thoracic neuropodia chaetal types. Naineris setosa only possesses capillaries, whereas N. aurantiaca has two types of uncini and subuluncini as well as capillaries. The shape of the furcate chaetae also differs between both these species: the tips of both tines are robust in N. aurantiaca , whereas in N. setosa they are longer and thinner. Another difference is that N. setosa does not possess branchial and interramal papillae, but N. aurantiaca does. Lastly, the proboscis in these two species differ; it is large and saclike in N. setosa but dendritic in N. aurantiaca . Naineris bicornis Hartman, 1951 was recorded in sediments from Ilha Grande, Rio de Janeiro ( Nonato 1981). This species differs from N. aurantiaca in that it lacks interramal papillae ( Amaral et al. 2013). There is only one record for N. bicornis from Brazil from Ilha Grande. Tellingly, the distinct shape of the prostomium in N. bicornis can also arise through the fixation methodology, as noted by Eisig (1914) in other species, so that sometimes the prostomium looks rounded and sometimes it looks spatulate. We requested material identified as N. bicornis from ZUEC-Pol, but the material listed in Amaral et al. (2013) is no longer available and as we cannot check this record, we suggest a misidentification of the specimen.

Figures

FIGURE 2. Naineris aurantiaca (neotype MNRJP 1956) A: Lateral view (Shirlastain-stained); B: Interramal papillae from abdominal chaetiger (Shirlastain-stained); C: Pre- and post-branchial papillae in abdominal segments (Shirlastain-stained); D: Chaetal distribution on thoracic segments (neuropodia). Legends: ip: interramal papilla, do: dorsal organ, bp: branchial papila, s: subuluncini; hu: heavily-ridged uncini; u: uncini; c: crenulated capillary (Scales: A: 500 µm; B: 200 µm; C: 100 µm; D: 50 µm).

FIGURE 3. Naineris aurantiaca from Marambaia Island (MNRJP 1957). A: Dorsal view of anterior region (Shirlastainstained); B: Lateral view of anterior region; C: Everted proboscis; D: Pygidium; E: Interramal papillae from abdominal chaetiger (Shirlastain-stained); F: Chaetal distribution on thoracic segments (neuropodia); G: Dorsal organs in abdominal segments (Shirlastain-stained); H: Pre- and post-branchial papillae in abdominal segments (Shirlastain-stained). Legends: ip: interramal papilla, do: dorsal organ, bp: branchial papilla (Scales: A-B: 2 mm; C: 1 mm; D-F-G: 200 µm; E: 100 µm; H: 500 µm).

FIGURE 4. Scanning electron microscopy of Naineris aurantiaca from Marambaia Island (MNRJP 1963). A: Dorsal view of anterior region; B: Surface of everted proboscis; C: Thoracic uncini; D: Chaetal arrangement in neuropodium; E: Abdominal spines in neuropodium; F: Furcate chaetae in abdominal notopodium. Legends: no: nuchal organ; s: subuluncini; hu: heavilyridged uncini; u: uncini; c: crenulated capillary (Scales A-B: 200 µm; C-F: 10 µm; D: 50 µm E: 5 µm).