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Asteroidea: Paxillosida) and Its Taxonomic Implications

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Asteroidea: Paxillosida) and Its Taxonomic Implications De los Palos-Peña, M., Solís-Marín, F.A., Laguarda- Figueras, A., & Durán-González. A. (2021). Ontogenetic variation of the odontophore of Luidia superba (Asteroidea: Paxillosida) and its taxonomic implications. Revista de Biología Tropical, 69(S1), 89-100. DOI 10.15517/rbt.v69iSuppl.1.46330 DOI 10.15517/rbt.v69iSuppl.1.46330 Ontogenetic variation of the odontophore of Luidia superba (Asteroidea: Paxillosida) and its taxonomic implications Magdalena De los Palos-Peña1*; https://orcid.org/0000-0002-9667-8670 Francisco Alonso Solís-Marín2; https://orcid.org/0000-0001-5729-3316 Alfredo Laguarda-Figueras2 Alicia Durán-González2 1. Posgrado en Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, México; [email protected] (*Correspondence). 2. Laboratorio de Sistemática y Ecología de Equinodermos, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, México; [email protected]; [email protected]; [email protected] Received 17-VI-2020. Corrected 14-IX-2020. Accepted 05-X-2020. ABSTRACT Introduction: The sea star odontophore is the structure positioned between the paired oral ossicles, with which they articulate through proximal and distal processes. The internal anatomy structures may be used as taxonomic characters for a precise differentiation between species, so it is necessary to describe the structures variation throughout growth. Objective: To describe the odontophore shape and variation of Luidia superba A. H. Clark, 1917 from specimens of the Gulf of California deposited in the Echinoderm National Collection, ICML UNAM. Methods: A total of 735 specimens were examined to describe the external characters, from which 55 selected specimens, within a range of R = 14 mm and R = 210 mm, were dissected to extract the odontophores and analyzed with geometric morphometrics. Results: Scanning electronic microscopy (SEM) images of the odontophores showing the variations in shape throughout growth are presented. Differences in shape between size groups were confirmed with a Canonical Variables Analysis (P < 0.05). Conclusions: There are three main groups in this size ranges where specialization of the stereom can be observed through the ontogenetic series; the variation in shape of the odontophore shown here is a precedent for the use of internal anatomy as new taxonomic characters of identification. Key words: Ossicles; morphology; oral frame; internal anatomy; geometric morphometrics; Scanning Electron Microscopy. The genus Luidia Forbes, 1839 is com- 5-11 quite long arms, which narrow distally posed of infaunal organisms; they usually (Downey, 1973; A.M. Clark & Downey, 1992; inhabit shallow waters and reefs on the conti- Benavides-Serrato, Borrero-Pérez, & Díaz- nental shelf, although some species can extend Sánchez, 2011). to the upper batial zone. They are generally Species of this genus are distributed in more active and agile than other starfish, and all oceans from the tropics to high temperate feed primarily on small mollusks and other latitudes. Some species have a wider distribu- echinoderms. They are characterized by having tion, while others have a more restricted one Revista de Biología Tropical, ISSN electrónico: 2215-2075, Vol. 69(S1): 89-100, March 2021 (Published Mar. 30, 2021) 89 (Sladen, 1889; A.M. Clark & Downey, 1992; variation that exists among specimens of the Puppin-Gonçalves, Rocha, Alencar, Moraes, Gulf of California to Peru. It is likely that a Araújo & Freire, 2020). The known morpho- successful predator, such as a luidid, needs a logical characteristics of the organisms that are developed oral system, since the five pairs of associated with the particulate nature of the interradial muscles, five radial pairs, the five substrate they inhabit are mainly the shape and transverse actinal muscles on the circumorals, arrangement of the ambulacral feet and paxil- and the five odontophore-oral muscles (Fig. 1) lae. These organisms are buried in the substrate, close the peristome by contracting. so it has been suggested that the rounded tip of The odontophore plays an important role the feet is adapted to push between the particles in the oral frame, since together with the of the substrate and contribute to an efficient oral ossicles, are responsible for the opening movement (Blake, 1989; Lawrence, 2013). movement of the mouth, which allows organ- Organisms of this genus are intraoral feed- isms feeding. Blake (1973) described that the ers, which may be related to their presence ontogenetic variation is greater than variation in particulate substrates where most of the between conspecific individuals of similar size, available food corresponds mainly to infaunal and that there is variation in age and size of the organisms, such as foraminifera. They do not various ossicles of an arm series, but he did not have the ability to separate the small infauna discuss this for the oral ossicles. It is necessary from the sediment while feeding; the car- to describe the variation of the odontophore diac stomach is very large, which gives them throughout its growth, especially in species that the ability to ingest very large prey (Hulings have a wide range of sizes such as this one. & Hemlay, 1963; Jangoux 1982a; Jangoux Studies on ossicle morphology are common 1982b; Lawrence, 2013). The mouth frame is in other echinoderm groups, such as ophiuroids composed of ten pairs of oral and circumoral (Thuy & Stöhr, 2016; Hendler, 2018; Alitto, muscles and five unpaired interradial odonto- Granadier, Christensen, O’Hara, Domenico & phores. This was described by Viguier in 1878, Borges, 2020). For asteroids, the taxonomic and he suggested that more precise characters usefulness of the individual elements and inter- may be derived from the oral frame and the pretations of evolutionary relationships have internal anatomy. The odontophore is made not been completely explored. The goal of this up of symmetrical, paired, distal, and proxi- study is to describe the ontogenetic variation of mal processes which articulate with the inner the odontophore shape of Luidia superba in the surface of the oral plates and an actinal keel Gulf of California from specimens deposited to which the fibers of the odontophore-oral in the Echinoderm National Collection, Insti- muscle attach (Gale, 2011). tuto de Ciencias del Mar y Limología, Univer- Luidia superba A.H. Clark, 1917 is the sidad Nacional Autónoma de México (ICML, largest species of the genus in the Pacific, and UNAM) as a first approach to evaluate the the largest specimen so far reported (R = 415 possibility of using it as a taxonomic character mm) was collected in Galapagos (Downey for differentiation at specific level. & Wellington, 1978) and deposited in the United States National Museum, Smithsonian MATERIALS AND METHODS Institution (USNM E18920). This species is distributed from the Gulf of California to Peru Specimen examination: We examined all (Alvarado & Solís-Marín, 2013). Although no the available specimens of L. superba depos- studies have been conducted on the feeding ited in the Echinoderm National Collection habits of this species, it is known that other ICML (UNAM) and the National Museum members of the genus are effective preda- of Natural History, Smithsonian Institution tors, so it is necessary to carry out studies on (USNM), Washington D. C. (735 specimens their anatomy and the intraspecific latitudinal in total). Organisms were observed using a 90 Revista de Biología Tropical, ISSN electrónico: 2215-2075 Vol. 69(S1): 89-100, March 2021 (Published Mar. 30, 2021) stereoscopic microscope Olympus SZX7 and rays being designated “B”, “C”, “D”, and “E” measured using a digital caliper (TRUPER clockwise when viewed from the oral side Caldi-6MP), the following measurements were (O’Neill, 1989; Lawrence, 2013). To standard- taken: R, r, arm length (LA) and arm width at the ize the odontophore selected per specimen, the base (BA). They were also photographed using pair of plates found in the interradium between a multifocal microscope LEICA Z16 APOA arms “C” and “D” (the arms between which the at the Laboratorio Nacional de Microscopía madreporite is found) was taken in each speci- y Fotografía de la Biodiversidad (Instituto de men. The dissection of the selected specimens Biología, UNAM). After reviewing the exter- was carried out using the method described nal characters of the available material to make by Fau and Villier (2018): specimens were the description, 55 of the Echinoderm National prepared in a dilute solution of NaClO (house- Collection (ICML-UNAM) specimens were hold bleach) followed by several rinsings with selected to represent the size range that this tap water and alcohol (70%); they were then species reaches in the Gulf of California; the dried, mounted and gold-coated on a SEM stub selected specimens range from R = 14 mm to at the Laboratorio Nacional de Microscopía R = 210 mm. We used the anatomical conven- y Fotografía de la Biodiversidad (Instituto de tions to describe ossicle orientation and nomen- Biología, UNAM) and at the National Museum clature described by Gale (2011) and Fau and of Natural History SEM Laboratory (USNM). Villier (2018): doda: distal oral/odontophore articulation on the oral; odom: odontophore- Geometric morphometrics: The SEM oral muscle; poda: proximal odontophore-oral images of the actinal view of the odontophore articulation (Fig. 1). were analyzed using geometric morphometrics with specialized software: TPSUtil, TPSDig, Scanning Electron Microscopy (SEM):
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