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Description of Scoloplos maranhensis sp. nov. (Orbiniidae, Annelida) from tropical Brazilian mangrove

Descrição de Scoloplos maranhensis sp. nov. (Orbiniidae, Annelida) do mangue tropical brasileiro

ABSTRACT

A new species of orbiniid, Scoloplos maranhensis sp. nov., is described from a tropical mangrove (Quebra Pote, state of Maranhão, Brazil, 02°41.344’S; 44°12.604’W) along with a comparative table with all Scoloplos species. The main distinguishing characters of S. maranhensis sp. nov. are thorax with 12 chaetigers, neuropodia from first chaetiger, notopodia and neuropodia without papillae, conical branchiae from 12th chaetiger to the end of the body, being larger and more robust in the anterior segments. Thoracic notopodia with seven crenulated capillary chaetae and abdominal notopodia with furcated and capillary chaetae. Thoracic neuropodia rounded with two crenulated capillary chaetae and six slender-smooth hooks, abdominal neuropodia with a post-chaetal lobe and only capillary chaetae. Papillae absent from posterior parapodia; one pair of cirriform anal cirri.

KEYWORDS
New species; Polychaeta; Biodiversity; South America

The family Orbiniidae is characterized by the prostomium shape, number of thoracic segments, localization of the first branchia, shape of the parapodia, the morphology and distribution of chaetae, and the body with two clearly distinct regions: thorax, which is dorsoventrally flattened and muscular, and the abdomen which is more cylindrical and delicate (Taylor, 1984Taylor, J. L. 1984. Family Orbiniidae Hartman, 1942. In: Uebelacker, J. M. & Johnson, P. G. eds. Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Vol. I. Vittor Mobile, Barry A, p. 1-38.). The worms of this family may have a long body with some adults of Orbinia reaching up to 300 mm, although other species do not exceed 4 mm in length such as those from the genus Proscoloplos (Taylor, 1984Taylor, J. L. 1984. Family Orbiniidae Hartman, 1942. In: Uebelacker, J. M. & Johnson, P. G. eds. Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Vol. I. Vittor Mobile, Barry A, p. 1-38.; Solís-Weiss et al., 2009Solís-Weiss, V.; Hermoso-Salazar, M.; Barbosa-López, A. & Hernández-Alcántara, P. 2009. Orbiniidae Hartman, 1942. In: de Léon-Gonzalez, J. A.; Bastida-Savala, L. F.; Carrera-Parra, M. E.; García-Garza, A.; Peña-Rivera, S. I.; Salazar-Vallejo, S. I. & Solís-Weiss, V. eds. Poliquetos (Annelida: Polychaeta) de Mexico y América Tropical. Monterrey, Universidad Autónoma de Nuevo Léon, p. 379-390.; Díaz et al., 2012Díaz, O. D.; Vanegas-Espinosa, V.; Cárdenas-Oliva, A. & Liñero-Arana, I. 2012. Orbiniidae Hartman, 1942 (Annelida: Polychaeta) de las costas de Venezuela. Biota Colombiana 13:3-19.). Orbiniids are excavators and deposit feeders, with a worldwide distribution occurring from intertidal to abyssal depths (Bleidorn, 2005 Bleidorn, C. 2005. Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data. Zoological Journal of the Linnean Society 144:59-73.; López et al., 2006López, E.; Cladera, P. & San Martin, G. 2006. Orbiniidae polychaetes (Polychaeta: Scolecida) from Coiba Island, eastern Pacific of Panama, with description of a new species. Revista de Biología Tropical 54:1307-1318.; Solís-Weiss et al., 2009Solís-Weiss, V.; Hermoso-Salazar, M.; Barbosa-López, A. & Hernández-Alcántara, P. 2009. Orbiniidae Hartman, 1942. In: de Léon-Gonzalez, J. A.; Bastida-Savala, L. F.; Carrera-Parra, M. E.; García-Garza, A.; Peña-Rivera, S. I.; Salazar-Vallejo, S. I. & Solís-Weiss, V. eds. Poliquetos (Annelida: Polychaeta) de Mexico y América Tropical. Monterrey, Universidad Autónoma de Nuevo Léon, p. 379-390.; Dean & Blake, 2015Dean, H. K. & Blake, J. A. 2015. The Orbiniidae (Annelida: Polychaeta) of Pacific Costa Rica. Zootaxa 3956:183-198.). Currently, two subfamilies are recognized Orbiniinae and Microrbiniinae (Díaz et al., 2012Díaz, O. D.; Vanegas-Espinosa, V.; Cárdenas-Oliva, A. & Liñero-Arana, I. 2012. Orbiniidae Hartman, 1942 (Annelida: Polychaeta) de las costas de Venezuela. Biota Colombiana 13:3-19.), encompassing 19 genera and more than 150 species, 26 of which have been recorded from Brazil (Blake, 2000Blake, J. A. 2000. A new genus and species of polychaete worm (Family Orbiniidae) from methane seeps. In: The Gulf of Mexico, with a review of the systematics and phylogenetic interrelationships of the genera of Orbiniidae. Cahiers de Biologie Marine 41(4):435-449.; Rouse & Pleijel, 2001Rouse, G. W. & Pleijel, F. 2001. Polychaetes. University Press, Oxford. 354p.; Leão & Santos, 2016Leão, L. S. D. & Santos, C. S. G. 2016. Orbinia (Polychaeta: Orbiniidae) from the Brazilian coast: two new species and two new records. Zootaxa 4105:145-158. ).

The genus Scoloplos was described by Blainville (1828Blainville, H. 1828. Dictionnaire des Sciences naturelles, dans lequel on traite méthodiquement des différens êtres de la nature, considérés soit en eux-mêmes, d’après l’état actuel de nos connoissances, soit relativement a l’utilité qu'en peuvent retirer la médicine, l’agriculture, le commerce et les arts. Suivi d’une biographie des plus célèbres naturalistes. FG Levrault, Strasbourg & Paris 57:1-628.) and belongs to the subfamily Orbiniinae. Scoloplos has traditionally been divided in two subgenera, Scoloplos (Scoloplos) and Scoloplos (Leodamas) (Hartman, 1957Hartman, O. 1957. Orbiniidae, Apistobranchidae, Paraonidae and Longosomidae. Allan Hancock Pacific Expeditions 15:211-393.; Pettibone, 1957Pettibone, M. H. 1957. North American genera of the family Orbiniidae. Journal of the Washington Academy of Sciences 47:159-167.; Day, 1967Day, J. H. 1967. A monograph on the Polychaeta of Southern Africa. London, British Museum (Natural History). 878p., 1973Day, J. H. 1973. New polychaeta from Beaufort, with a key to all species recorded from North Carolina. NOAA Technical Report Series National Marine Fisheries Service Circular 375:1-140.; Fauchald, 1977Fauchald, K. 1977. The polychaete worms, definitions and keys to the orders, families, and genera. Natural History Museum of Los Angeles County Science ; Eibye-Jacobsen, 2002Eibye-Jacobsen, D. 2002. The Orbiniidae (Annelida: Polychaeta) of the BIOSHELF Project, Andaman Sea, Thailand . Phuket Marine Biological Center, Special Publication 24:77-99.). More recently both have been considered two separated genera, although the phylogeny of the group is not completely resolved yet (Blake, 2000Blake, J. A. 2000. A new genus and species of polychaete worm (Family Orbiniidae) from methane seeps. In: The Gulf of Mexico, with a review of the systematics and phylogenetic interrelationships of the genera of Orbiniidae. Cahiers de Biologie Marine 41(4):435-449., 2017Blake, J. A. 2017. Polychaeta Orbiniidae from Antarctica, the Southern Ocean, the Abyssal Pacific Ocean, and off South America. Zootaxa 4218:1-145.; Bleidorn et al., 2009Bleidorn, C., Hill, N., Erséus, C. & Tiedemann, R. (2009) On the role of character loss in orbiniid phylogeny (Annelida): molecules vs. morphology. Molecular Phylogenetics and Evolution 52: 57-69. http://dx.doi.org/10.1016/j.ympev.2009.03.022.
http://dx.doi.org/10.1016/j.ympev.2009.0...
; Zhadan et al., 2015Zhadan, A.; Stupnikova, A. & Neretina, T. 2015. Orbiniidae (Annelida: Errantia) from Lizard Island, Great Barrier Reef, Australia with notes on orbiniid phylogeny. Zootaxa 4019 (1):773-801. http://dx.doi.org/10.11646/zootaxa.4019.1.27
http://dx.doi.org/10.11646/zootaxa.4019....
; Read & Fauchald, 2018Read, G. & Fauchald, K. 2018. World Polychaeta database. Orbiniidae Hartman, 1942. Available at <Available at http://www.marinespecies.org/aphia.php?p=taxdetails&id=902 >. Accessed on 04 March 2018.
http://www.marinespecies.org/aphia.php?p...
), classification followed here. Scoloplos has four or less foot papillae plus stomach papillae, thoracic neuropodial hooks accompanied with capillary chaetae, and branchiae starting from chaetiger 8 or later, while Leodamas have numerous hooks in thoracic neuropodia accompanied with few or no capillary chaetae, single thick projecting aciculae in abdominal neuropodia, and branchiae starting before chaetiger 7 (Bleidorn et al., 2009Bleidorn, C.; Kruse, I.; Albrecht, S. & Bartolomaeus, T. 2005. Mitochondrial sequence data expose the putative cosmopolitan polychaete Scoloplos armiger (Annelida, Orbiniidae) as a species complex. BCM Evolutionary Biology 6:1-13. http://dx.doi.org/10.1186/1471-2148-6-47.
http://dx.doi.org/10.1186/1471-2148-6-47...
; Zhadan et al., 2015Zhadan, A.; Stupnikova, A. & Neretina, T. 2015. Orbiniidae (Annelida: Errantia) from Lizard Island, Great Barrier Reef, Australia with notes on orbiniid phylogeny. Zootaxa 4019 (1):773-801. http://dx.doi.org/10.11646/zootaxa.4019.1.27
http://dx.doi.org/10.11646/zootaxa.4019....
). Currently, there are 29 species of Scoloplos described, seven of which already recorded from Brazilian coast (Amaral et al., 2012Amaral, A. C. Z.; Nallin, S. A. H. & Steiner, T. M. 2012. Catálogo das espécies dos Annelida Polychaeta do Brasil. Unicamp, Campinas. Available at <Available at http://www.ib.unicamp.br/destaque/jbiota/bentos_marinho/prod_cient/texto_poli.pdf >. Accessed on 15 March 2015.
http://www.ib.unicamp.br/destaque/jbiota...
; Read & Bellan, 2017Read, G. & Bellan, G. 2017. Scoloplos Blainville, 1828. In: Read, G. & Fauchald, K. eds. World Polychaeta database. Available at <Available at http://www.marinespecies.org/aphia.php?p=taxdetails&id=129425n >. Accessed on 05 February 2018.
http://www.marinespecies.org/aphia.php?p...
). In the present study, we describe an additional species, Scoloplos maranhensis sp. nov. from tropical Brazilian mangroves and provide a comparative table of all species of the genus.

MATERIAL AND METHODS

All material studied was sampled at Quebra Pote mangrove, located in the city of São Luís, Maranhão, Brazil (02°41.344’S; 44°12.604’W; Fig. 1). The mangrove is strongly influenced by tides, with amplitude of up to 8 m (Silva & Almeida, 2002Silva, J. R. R. & Almeida, Z. S. 2002. Zoneamento vertical dos crustáceos bentônicos em substratos inconsolidados do manguezal do Quebra-Pote na ilha de São Luís, Maranhão - Brasil. Boletim Técnico-Científico CEPENE 10:125-143.), and by the direct interference of domestic sewage discharged in natura. Four sampling campaigns were performed in November 2013, January 2014, March 2014, and July 2014 using a corer with 10 cm diameter introduced 20 cm depth in the sediment. On each occasion samples were taken in the lower, intermediate and upper mesolittoral. The material sampled was fixed in 4% formalin, and in laboratory washed in 0.5 mm pore diameter sieves and preserved in 70% ethanol.

Fig. 1
Type locality of Scoloplos maranhensis sp. nov., Quebra Pote Mangrove, Maranhão, Brazil.

Descriptions were based on external morphology, mostly from anterior and median part of the body where most of the diagnostic characters are found. Descriptions followed the classical taxonomic style, following the terminology adopted by Leão & Santos (2016Leão, L. S. D. & Santos, C. S. G. 2016. Orbinia (Polychaeta: Orbiniidae) from the Brazilian coast: two new species and two new records. Zootaxa 4105:145-158. ). In addition, the description included body length, always relating to the number of segments. The median width was measured including the parapodia but excluding the chaeta. To better visualization of the papillae, the Shirlastain A® dye was used in some individuals. Type material was deposited at the “Coleção de Invertebrados Paulo Young” (CIPY) from Universidade Federal da Paraíba.

RESULTS AND DISCUSSION

Scoloplos maranhensis sp. nov.

urn:lsid:zoobank.org:act:B2DA4CC4-26D5-483A-9CFC-44718328E A2F

(Figs 2-12)

Figs 2-4
Scoloplos maranhensis sp. nov., holotype: 2, general view; 3, 4, detail of the anterior region with the prostomium and the smooth proboscis. Scales: Fig. 2: 2 mm; Figs 3, 4: 0.5 mm.

Figs 5-8
Scoloplos maranhensis sp. nov., anterior view of parapodium of the paratype: 5, parapodium 7, capillary notopodial chaetae and crenulate neuropodial chaetae; 6, Parapodium 7, notopodial lobe; 7, parapodium 18; 8, parapodium 30, furcate chaetae. Scales: Figs 5, 6: 0.1 mm; Fig. 7: 0.2 mm; Fig. 8: 0.05 mm.

Figs 9-12
Scoloplos maranhensis sp. nov.: 9, crenulate chaetae; 10, furcate chaetae (black arrow); 11, detail the smooth hooks; 12, detail of posterior region with the pygidial cirri. Scales: Figs. 9, 10: 0.1 mm; Fig. 11: 0.01 mm; Fig. 12: 0.5 mm.

Type material. Holotype, Mangrove from Quebra Pote, São Luís, state of Maranhão, Brazil (02°41.344’S; 44°12.604’W), 09 January 2014 (CIPY - 1730).

Paratypes. A total of 11 paratypes, length ranging for 1.4 to 0.2 mm and number of segments from 10 to 80. Quebra Pote mangrove, Brazil (02°41.344’S; 044°12.604’W): 05 November 2013, (1 paratype, CIPY - 1731); 07 January 2014; (2 paratypes, CIPY - 1732); 03 September 2014 (6 paratypes, CIPY 1733); 28 September 2012 (2 paratypes, CIPY 1734).

Diagnosis. Thorax with 12 segments, chaetae appearing since the first segment. Notopodia and neuropodia without papillae. Conical branchiae from chaetiger 12 to the end of the body. Branchiae larger and more robust in the anterior segments. Each branchial pair located on median dorsal part. Cirriform post-chaetal notopodial lobes, inconspicuous at first, but long from chaetiger 3. Thoracic notopodia with seven crenulated capillary chaetae, neuropodia with two crenulated capillaries and six slender-smooth hooks. Abdominal notopodia with crenulated capillary and furcate chaetae and and neuropodia crenulated capillary chaetae.

Description. The holotype is a complete specimen with 57 segments, 3.0 mm length, 0.1 mm wide. Body long, very slender and tapered at the posterior end (Fig. 2). Prostomium conical, longer than wide (Fig. 3). Eyes, antennae and tentacular cirri absent, nuchal organs present but not visible (Fig. 3). Peristomium as a single achaetous ring, similar in length to segment 1. Undivided smooth proboscis covered with diffusely distributed rounded small papillae (Figs 3, 4). Thorax dorsoventrally flattened with 12 chaetigers, widest in the middle part. Slight transition from thorax to abdomen from segment 12. All parapodia birramous, present from the first chaetiger. Abdominal neuropodia rounded, notopodia with one dorsal lamella and all ventral lamella absent. Thoracic notopodia and neuropodia without papillae. Branchiae conical, on median-dorsal position, beginning from the 12th segment and continuing through the body. Branchiae at first long and robust, becoming thinner towards the posterior end. Cirriform post-chaetal notopodial lobes inconspicuous at first, but long from chaetiger 3; neuropodia rounded on thoracic chaetigers and with a developed lobe in abdominal chaetigers. Thoracic notopodia with crenulated capillaries (Figs 5, 6, 9, 10) and abdominal notopodia with crenulated capillaries and furcated chaetae with unequal ends and internally hairy (Figs 7, 8, 10). Thoracic neuropodia rounded with two crenulated capillaries chaetae, one single neuropodial acicular spine in the segments and six slender, smooth hooks (Figs 5, 6, 11); abdominal neuropodia with post-chaetal lobe and only crenulated capillaries (Fig. 7). Anal papillae absent. Pygidium with one pair of cirriform anal cirri (Fig. 12).

Color. Preserved specimens are yellowish opaque.

Remarks. Scoloplos maranhensis sp. nov. can be clearly placed in the genus Scoloplos due to the position of the branchiae and the presence of thoracic neuropodial hooks accompanied with capillary chaetae (Blake, 2000Blake, J. A. 2000. A new genus and species of polychaete worm (Family Orbiniidae) from methane seeps. In: The Gulf of Mexico, with a review of the systematics and phylogenetic interrelationships of the genera of Orbiniidae. Cahiers de Biologie Marine 41(4):435-449.; Zhadan et al., 2015Zhadan, A.; Stupnikova, A. & Neretina, T. 2015. Orbiniidae (Annelida: Errantia) from Lizard Island, Great Barrier Reef, Australia with notes on orbiniid phylogeny. Zootaxa 4019 (1):773-801. http://dx.doi.org/10.11646/zootaxa.4019.1.27
http://dx.doi.org/10.11646/zootaxa.4019....
). The main distinctive characters of S. maranhensis sp. nov. is the thoracic notopodia with seven crenulated capillary chaetae, neuropodia with two crenulated capillaries and six slender-smooth hooks. Only four species may have 12 thoracic segments with branchiae first appearing on the 12th segment: Scoloplos texanaMaciolek & Holland, 1978Maciolek, N. J. & Holland, J. S. 1978. Scoloplos texana. A new orbiniid polychaete from South Texas, with notes on the related species Scoloplos treadwelli Eisig. Contributions in Marine Science 21:161-169. from Gulf of Mexico, Scoloplos intermedius (Hartman, 1965Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas. Occasional Papers of the Allan Hancock Foundation. 28:1-384.) from New England, Scoloplos armiger (Müller, 1776) from Denmark and Scoloplos suroestense Blake, 2017 from Punta Suroestense, Chile Bay (Tab. I). However, in all these three species both characters are variable (Hartman, 1965Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas. Occasional Papers of the Allan Hancock Foundation. 28:1-384.; Maciolek & Holland, 1978Maciolek, N. J. & Holland, J. S. 1978. Scoloplos texana. A new orbiniid polychaete from South Texas, with notes on the related species Scoloplos treadwelli Eisig. Contributions in Marine Science 21:161-169.; Mackie, 1987Mackie, A. 1987. A review of species currently assigned to the genus Leitoscoloplos Day, 1977 (Polychaeta: Orbiniidae), with descriptions of species newly referred to Scoloplos Blainville, 1828. Sarsia 72:1-28.; Blake, 2017Blake, J. A. 2017. Polychaeta Orbiniidae from Antarctica, the Southern Ocean, the Abyssal Pacific Ocean, and off South America. Zootaxa 4218:1-145.), while in all the 12 specimens of S. maranhensis sp. nov. examined here the thorax had exactly 12 chaetigers long and the branchiae always first appeared in this last thoracic segment (Tab. I). Scoloplos armiger also always have 12 thoracic segments (Uebelacker & Johnson, 1984Uebelacker, J. M. & Johnson, P. G. 1984. Taxonomic guide to the polychaetes of the northern Gulf of Mexico. Louisiana, Minerals Manegement Service, U.S Departement of Interior , p.1-38.) likewise S. maranhensis sp. nov., however they differ in the position of the first branchiae, in the 10th and 12th segment respectively (Tab. I). Other characteristics differing these species from S. maranhensis sp. nov. include: i) the prostomium shape, conical in S. maranhensis sp. nov. and pointed and/or triangular in the others; ii) the absence of anal cirri in S. suroestense; iii) the presence of papillae in S. armiger and S. intermemdius; and iv) the chaetal arrangement of S. texana including, for instance, serrated hooks in the neuropodia from thoracic chaetae which are smooth in S. maranhensis sp. nov. (Tab. I and references therein).

Tab. I
Main characters of currently known Scoloplos species.

Etymology. The species was named in reference to Maranhão, the State with the largest coastline from Brazil form where S. maranhensis sp. nov. was found.

Habitat. Muddy mangrove substrate with temperature between 25.7 and 27.8°C and salinity between 11 and 30.

Geographical distribution. Atlantic Ocean, Brazil, Quebra Pote Mangrove, São Luís, Maranhão State (02°41.344’S; 044°12.604’W).

Acknowledgements

We thank the Brazilian National Research Council CAPES. Universidade Estadual do Maranhão (UEMA), and the Graduate Program in Recursos Aquáticos e Pesca (UEMA). The group Annelida for collaborating in the sending of old references. Dr. Vasily Radashevsky of the National Scientific Center of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences helped translating old descriptions written in Russian. Dr. Hilton Tulio Costi of the Institutional Laboratory of Electronic Microscopy of the Museu Paraense Emílio Goeldi.

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Publication Dates

  • Publication in this collection
    25 Feb 2019
  • Date of issue
    2019

History

  • Received
    07 July 2018
  • Accepted
    29 Jan 2019
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