Diversity of fish otoliths from the Gulf of Mexico and Caribbean Sea: report on the first digital collection of fish otoliths from the Atlantic region of Mexico

15 The Otolith Collection of Fishes from the Gulf of Mexico and Caribbean Sea was created 16 with the objective of conserving and illustrate the diversity of these structures from 17 species living in the Gulf of Mexico, the Caribbean Sea, and the freshwater and brackish 18 systems of the Yucatan Peninsula, incorporating morphological descriptions and 19 morphometric data. Otoliths, non-skeletal calcareous structures that develop in the inner 20 ear of fish, are essential for balance and hearing. They have become pivotal tools for age 21 and growth determination, population analysis, and ecological, trophic, and archaeological 22 studies due to their resistance to degradation and the extensive information they provide 23 about the environment and physiology of fish. The collection now includes otoliths from 24 214 species from 67 families, obtained through sampling campaigns and collaborations 25 with local fishermen. The otoliths are extracted using techniques that ensure the integrity 26 of the structures for later preservation. High-resolution images are obtained using optical 27 and scanning electron microscopy, and these images are stored in an online database. 28 This database facilitates research and teaching by providing public access to digital 29 specimens and associated data. In addition to fostering academic development, this 30 collection represents a significant step towards the creation of a national platform for 31 otolith data analysis, aligned with international efforts to digitize biological collections. 32 Despite the financial and logistical challenges involved in building and maintaining 33 biological collections, this collection demonstrates its value as an essential resource for 34 taxonomic, systematic, and ecological studies, as well as for biodiversity education and 35 awareness. The online availability of the collection not only facilitates access to data but 36 also promotes innovation and international collaboration in the study of fishes and their 37 habitats. 38 39

Biological collection, digitization, fish, image analysis, online collection 40 41

42 Throughout history, biological collections have played a fundamental role in documenting 43 the biodiversity of our planet. They provide essential information for the knowledge, 44 exploitation, and sustainable use of our biological capital, and they assist in the analysis 45 of trends derived from environmental changes. Additionally, they contribute necessary 46 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 information for studies on environmental health, epidemiology, and national security, 47 proving to be irreplaceable tools for investigating cases of biological terrorism (Suarez 48 and Tsutsui 2004) and determining the effects of anthropogenic activities (Izzo et al. 49 2018). The information generated from the analysis of specimens kept in biological 50 collections has motivated changes in public policies on the sustainable use of natural 51 resources, significantly impacting social welfare (National Academies of Sciences, 52 Engineering, and Medicine 2020). 53 54 Biological collections are not only essential repositories for taxonomic and systematic 55 work on the biota but also contribute to the development of other scientific disciplines. 56 They play a crucial role in education by raising awareness of the need to document and 57 conserve biodiversity and by enabling the development of new skills in research and data 58 analysis. Therefore, the role of biological collections transcends purely scientific 59

and should be considered necessary tools that benefit society (Kellner 2024). 60 61 At different regional levels, collections maintain and catalog specimens of local 62 ecosystems, creating, preserving, and increasing information on species distribution, 63 identifying endemic or invasive species, and evaluating changes in biodiversity over time 64 (Meineke et al. 2018). Thus, biological collections are essential references in conservation 65 efforts, including those focused on endangered species and their habitats. 66 67 As fishes have been a group of interest for centuries, they are part of several biological 68 collections, notable for their diversity of forms and adaptations. Approximately 36893 fish 69 species are recognized (Fricke et al. 2024), equivalent to more than half of all living 70 vertebrates. Mexico, with its high diversity of fishes, boasts at least 2763 species 71 (Espinosa-Pérez 2014), of which 1816 are found in the Gulf of Mexico and the Caribbean 72 Sea (Robertson and Van Tassel, 2023). In this region, fishing activities by industrial and 73 artisanal fleets, along with recreational activities such as diving and sport fishing, are 74 depleting natural populations (De la Cruz et al. 2016). Given that many fish species 75 present on the Atlantic coast of Mexico have economic, nutritional, cultural, ecological, or 76 evolutionary importance, it is necessary to know the specific identity of organisms and 77 study aspects of their biology, ecology, and biogeography, underscoring the relevance of 78 scientific collections. 79 80 Although specimens deposited in ichthyological collections are useful for various research 81 purposes, their importance in paleontological, archaeological, and trophic studies is 82 particularly noteworthy. In trophic studies, for example, prey of piscivorous organisms are 83 often identified by examining stomach contents, vomit, or feces (Baker et al. 2024). This 84 methodology is limiting when the prey's degree of digestion is advanced. However, other 85 structures such as otoliths (Baker et al. 2014), which may be present in the digestive tract 86 of ichthyophagous organisms due to their resistance to degradation and retention of 87 original characteristics in many situations (Rivaton and Bourret 1999; Tuset et al. 2008), 88 can be recognized. In paleontology and archaeology, zooarchaeological remains, 89 including otoliths, are valuable for reconstructing environmental conditions, such as the 90 distribution and extent of water bodies, and recognizing the species used by ancient 91 cultures (Lin et al. 2022). 92 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 93 Otoliths are non-skeletal calcareous concretions (Nolf 1985; Maisey 1987) that grow 94 within the vestibular system of vertebrates, particularly in the membranous labyrinth of the 95 inner ear. In bony fish, the inner ear consists of three semicircular canals and three otic 96 chambers, each housing an otolith. Depending on the chamber in which they are located, 97 otoliths are called sagitta (in the sacculum), lapillus (in the utricle), and asteriscus (in the 98 lagena). These structures play a crucial role in balance, orientation, and sound detection 99 (Platt and Popper 1981; Gauldie and Nelson 1988; Campana and Thorrold 2001). Otoliths 100 are composed of calcium carbonate (CaCO3) in the form of aragonite and other inorganic 101 salts, with crystals embedded in an organic matrix composed of a fibrous protein called 102 otolin, deposited concentrically around a nucleus (Martínez-Pérez et al. 2018). The 103 presence of several elements is related to metabolically regulated processes and is 104 influenced by both endogenous and exogenous factors. Therefore, otoliths can provide 105 information on the physiological aspects of organisms and the environments in which they 106 developed (Volpedo and Echeverría 2003). These characteristics have made otoliths key 107 tools in research on age and growth (Francis and Campana 2004), population dynamics, 108 trophic studies, archaeological analyses, and present and past environmental conditions. 109 110 Sagittae have a particular morphology that varies among species, making them useful as 111 auxiliary structures in organism identification and for distinguishing between 112 phylogenetically close species (Volpedo and Echeverría 2003). Analyzing the shape, size, 113 and structural characteristics of otoliths not only allows for species identification but also 114 helps investigate evolutionary relationships and reconstruct phylogenetic histories. 115 116 Rojo (2015) emphasized the importance of incorporating otoliths into fish collections, as 117 they enable the extraction of biological information about species that is otherwise 118 unattainable through the analysis of other structures such as spines, scales, or bones. 119 Otoliths store unique data on the life history of organisms and the environmental 120 conditions of their habitats. It is in this context that the Otolith Collection of Fishes from 121 the Gulf of Mexico and Caribbean Sea was created and registered with the Mexican 122 Ministry of Environment and Natural Resources (SEMARNAT, registration code DGVS-123 CC-305-18). 124 125 The main objective of this collection is to preserve otoliths of a wide variety of species, 126 including those of economic and ecological importance, as well as endangered and 127 endemic species from the Gulf of Mexico and Caribbean Sea, and freshwater and 128 brackish systems of the Yucatan Peninsula. The addition of the Mexican Atlantic Fish 129 Otolith Collection (Martínez-Pérez et al. 2011; Del Moral-Flores et al. 2016) and proper 130 curation and maintenance have turned this collection into a national reference, 131 contributing to research in disciplines such as biology, ecology, biogeography, 132 archaeology, and fisheries, among others. To enhance this purpose and facilitate access 133 to specimens and other resources such as databases or software for shape analysis, this 134 collection is available online. Public access to information is thus guaranteed, scientific 135 collaboration is encouraged, and educational and training opportunities are improved 136 (Monfils et al. 2017), extending these benefits to a wide geographical extent. 137 138 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421

139 Specimen collection 140 Since 2011, multiple campaigns have been conducted in various localities of the Gulf of 141 Mexico and the Caribbean Sea to obtain bony fish specimens, which serve as the primary 142 input of biological material. Specific sampling methods have been employed, involving 143 different fishing gears operated under catch permits obtained from the competent 144 authorities. Additionally, visits have been made to markets, fish markets, and ports, and 145 agreements have been established with fishermen from different areas to direct sampling 146 efforts towards certain species of interest. The Otolith Collection of Fishes from the Gulf of 147 Mexico and Caribbean Sea has been enriched by the addition of the Mexican Atlantic 148 Fish Otolith Collection (Martínez-Pérez et al., 2011; Del Moral-Flores et al., 2016). 149 150 It is important to note that otolith extraction requires that the specimens be fresh or frozen, 151 as otoliths preserved in unneutralized formalin or alcohol degrade, reducing the chances 152 of correct identification (Hecht, 1990) and limiting their usefulness as collection 153 specimens. 154 155 After collection, each fish is identified to the species level using taxonomic keys and 156 guides. Biometric data, such as total length (TL), standard length (SL), cephalic length 157 (CL), body height, weight, and, when possible, sex, are recorded for each specimen. 158 Capture data, including locality, date, and geographic coordinates, are also documented. 159 160 Otolith extraction 161 Two techniques are employed for otolith extraction. One method involves lifting the 162 operculum, removing the gills, and breaking the bony capsules to expose and extract the 163 otoliths with forceps. The other method involves separating the head from the body, 164 removing the gill arches and surrounding tissue to locate the bony capsules. The blades 165 of a pair of scissors are inserted into the center of the first vertebra, using them as a lever 166 to lift the ventral bone of the capsule and expose the otoliths. These procedures are 167 performed under a stereoscopic microscope. 168 169 Once extracted, the otoliths are cleaned of residual tissue, and detailed morphometric 170 measurements, including perimeter, area, height, width, and acoustic sulcus dimensions, 171 are obtained using Zen Pro software (Zeiss). These data allow for the calculation of 172 different morphometric indices useful in species comparison. 173 174 The terminology used for the morphological descriptions of sagittae is derived from the 175 works of Volpedo and Echeverría (2000), Mascareñas-Osorio et al. (2003), Tuset et al. 176 (2008), and Martínez-Pérez et al. (2018). 177 178 Otoliths are stored in vials labeled with biometric and capture data, including a unique 179 identification code for each specimen. The vials are placed in boxes, which are labeled 180 and stored in the collection under controlled temperature and humidity conditions to 181 ensure long-term preservation. 182 183 Digitization of the collection 184 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 The digitization of the Otolith Collection of Fishes from the Gulf of Mexico and Caribbean 185 Sea is a key component in improving the accessibility and usefulness of the stored 186 specimens. This collection includes a photographic archive of approximately 2600 otolith 187 images obtained using optical and scanning electron microscopy techniques. These 188 images are used not only for documentation and preservation but also to facilitate 189 research and comparative analysis. 190 191 High-resolution images of the otoliths are obtained using an advanced Zeiss AxioZoom 192 stereo microscope, which provides continuous magnification and excellent depth of field, 193 crucial for capturing the morphological details of the structures. These images are used 194 for both taxonomic identification and morphometric studies. Additionally, a scanning 195 electron microscope (Jeol 7600F FESEM) is used to obtain high-resolution images with a 196 large depth of field, allowing for the visualization of ultrastructural details of otoliths not 197 perceptible with optical microscopy. This equipment is essential for studies requiring 198 detailed analysis of the otolith surface and its microstructures. 199 200 Images are selected to highlight the distinctive characteristics of each otolith, with angles 201 and focus optimized for this purpose. Each image is edited to standardize the black 202 background, enhancing contrast and facilitating the visualization of morphological details. 203 A watermark with the collection logo is added to each image to ensure provenance and 204 maintain the collection's identity. 205 206 The digitized images are stored in a database management system that allows easy 207 access and retrieval. Each image is associated with the biometric and capture data of the 208 corresponding specimen, including the unique identification code. 209 210

211 Physical specimens 212 The Otolith Collection of the Gulf of Mexico and Caribbean Sea comprises 3001 lots, 213 representing 214 species from 67 families (Table 1). Sciaenidae is the most represented 214 with 17 species, followed by Carangidae and Haemulidae, with 15 and 11 species, 215 respectively. 216 217 The species with the highest number of specimens in the collection are Menticirrhus 218 americanus (Linnaeus, 1758) with 191 lots, Ariopsis felis (Linnaeus, 1766) with 108 lots, 219 Bagre marinus (Mitchill, 1815) with 106 lots, and Haemulon plumierii (Lacepède, 1801) 220 with 102 lots. Each lot typically contains all three pairs of otoliths: sagitta, asteriscus, and 221 lapillus. 222 223 The collection includes otoliths from specimens collected along the coasts of the states of 224 Tamaulipas, Veracruz, Campeche, and Yucatan, which is the best-represented state with 225 1700 lots, followed by Campeche with 894 lots and Veracruz with 368 lots. Additionally, 226 the collection contains 23 lots of otoliths from Cuba. 227 228 Fig. 1 illustrates the sagittal otoliths of most of the families included in the collection. 229 These images on the collection's web page are accompanied by morphological 230 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 descriptions and morphometric characteristics such as length, height, perimeter, and area 231 of both the sagitta and the acoustic sulcus. These variables allow for the calculation of 232 shape indices, including circularity, rectangularity, aspect ratio, and the proportion of the 233 area occupied by the acoustic sulcus. The diversity of the shape of the saccular otoliths is 234 notable, ranging from simple ellipsoids to complex patterns with species-specific 235 projections and invaginations. The dorsal and ventral margins can be smooth, scalloped, 236 lobed, serrated, or serrate. The anterior and posterior regions exhibit significant variability, 237 being pointed, angulated, rounded, truncate, oblique, lanceolate, bilobed, double-pointed, 238 or irregular. Many species have an indentation in the anterior region, the ostial fissure, 239 which delineates the rostral and antirostral regions. The characteristics of the fissure, 240 whether deep or shallow, angulated or rounded, affect the shape and size of these 241 sections. The acoustic sulcus, a longitudinal depression along the medial face of the 242 sagitta, consists of the ostium anteriorly and the cauda posteriorly. In some instances, 243 thickening of the edges, known as cristae, and dorsal or ventral depressions are present 244 along the acoustic sulcus. 245 246 Website 247 The online catalog of the Fish Otolith Collection of the Gulf of Mexico and Caribbean Sea 248 aims to organize and present the information in a user-friendly yet comprehensive 249 manner, available to academics, students, and anyone interested in fish biology and 250 ecology. 251 252 The website features a detailed database with information on each otolith, including 253 species, location, size and weight of the specimen, collector, and morphological 254 parameters of the otolith. It also includes a gallery of images or scientific illustrations of 255 the species and photographs of the otoliths obtained through optical and electron 256 microscopy. Detailed morphological descriptions of the left sagitta of each species are 257 accessible. 258 259 Before publication on the website, the quality of the digital data is verified to ensure high-260 value information that can positively impact research and education. 261 262 To facilitate consultation, the webpage offers search functions by taxon (family, genus, or 263 species) and keywords related to otolith characteristics. Additionally, didactic resources 264 such as videos on otolith extraction and description techniques, and the use of relevant 265 equipment for their study are available. The otolith collection webpage can be accessed 266 at https://otolitos.unam.mx. 267 268 Publications and analytical tools 269 Derived from the curatorial work of the collection and the information on each specimen, a 270 catalog with descriptive cards of the sagittal otoliths of 155 fish species from the Gulf of 271 Mexico was published (Martínez-Pérez et al., 2018). This catalog includes standardized 272 descriptions of the shape, as well as some morphometric data of the structure, acoustic 273 sulcus, and shape indices. Currently, a second edition of the catalog is being prepared, 274 which will include information on species added in recent years. 275 276 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Additionally, a software, Invariant Otolith Shape Analysis (IOSA), which can be 277 downloaded here, https://sites.google.com/view/jorge-perez/community#h.hh1lv19s6hlj ) 278 has been developed to automatically obtain morphometric descriptors and shape 279 invariants of otoliths from their images, based on the methodology proposed by Hevia-280 Montiel et al. (2021). 281 282 Recently, the collection has been used to corroborate the presence of fish remains in the 283 Mayan archaeological record. This information will deepen our understanding of pre-284 Hispanic fishing practices and infer possible changes in some fish populations in the 285 region. 286 287

288 The Otolith Collection of the Gulf of Mexico and Caribbean Sea is a sample of the 289 richness and ichthyological diversity of the region. It has a collection of 214 species 290 belonging to 67 families, which represent a significant percentage (just over 11%) of the 291 total number of species recorded for the Gulf of Mexico and the Caribbean Sea. 292 (Robertson and Van Tassel, 2023). The families and species most represented in the 293 collection reflect the specific composition of the local ichthyofauna, its ecological 294 importance, and its relevance to the regional economy. This extensive taxonomic 295 representation provides an invaluable database for biodiversity, taxonomy, and 296 systematic studies of fishes in the region. 297 298 The otoliths in this collection come from specimens obtained in various geographic 299 regions, enabling the study of species distribution and population variability in different 300 localities of the Gulf of Mexico and the Caribbean. With 3001 lots in total, the collection 301 provides a robust data set for statistical analyses, intraspecific variability studies, and 302 temporal trend evaluations. 303 304 Otoliths are invaluable in paleontological, archaeological, and biological explorations, 305 such as analyzing the stomach contents of piscivorous species. Reference collections 306 have been established to assist in species identification from remains found in 307 archaeological contexts, as documented by Disspain et al. (2016) and Lambrides et al. 308 (2024). The catalogs by Mascareñas et al. (2003) and Martínez-Pérez et al. (2018), along 309 with the website www.otolitos.unam.mx, support research efforts in Mexico. 310 311 The digitization of biological collections, including the Otolith Collection of the Gulf of 312 Mexico and Caribbean Sea, is crucial for making scientific information universally 313 accessible. Digitization transforms specimen information into digital formats, allowing for 314 easy access and analysis by anyone with an internet connection. This approach 315 modernizes the use of biological collections, eliminating the need for physical travel or 316 loan requests, which risk loss or damage to the specimens. 317 318 Digitization also fosters new research topics and addresses global challenges related to 319 climate change, food security, and conservation (Soltis, 2017). The online presence of the 320 Otolith Collection makes it the first digital otolith collection in Mexico, facilitating 321 international collaboration and knowledge dissemination. This effort aligns with global 322 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 initiatives like the "Integrated Digitized Biocollections" (iDigBio), the "European Distributed 323 System of Scientific Collections" (DiSSCo), and the "Innovation and Consolidation for 324 Large Scale Digitization of Natural Heritage" (ICEDIG). These programs promote the 325 networking of collections worldwide, contributing to the "global museum" concept (Bakker 326 et al., 2020). 327 328 Despite the benefits of digitization, physical specimens remain the primary source of 329 verifiable data. Digital information complements but does not replace physical specimens. 330 Increased online accessibility often leads to more visits to collections and loan requests 331 for physical specimens (Vollmar et al., 2010). Therefore, preserving physical specimens in 332 optimal conditions is a primary task for the curatorial staff of the Otolith Collection. 333 334 Additionally, the webpage offers accessible image processing software that employs 335 invariant descriptors, as detailed by Hevia-Montiel et al. (2021). These descriptors are 336 independent of rotation, scale, or translation, thereby ensuring consistent results. This 337 software provides additional information beyond traditional descriptors, such as otolith 338 area or perimeter (Hevia-Montiel et al., 2021), which is particularly beneficial in taxonomic 339 studies that rely on limited structures for species identification. By offering a user-friendly 340 interface that allows the application of image processing through a webpage, the tool 341 significantly broadens its user base. Enhancing collaboration is crucial, especially in 342 studies involving these structures, where there is an increasing demand for web-based 343 tools that enable researchers to browse and search images and data in conjunction with 344 biological analyses. 345 346 The contributions of biological collections to knowledge, education, and innovation are 347 immense but often unrecognized (National Academies of Sciences, Engineering, and 348 Medicine, 2020; Kellner, 2024). Many scientific communities have highlighted the risks of 349 inadequate funding for maintaining the infrastructure necessary for the proper functioning 350 of these collections (Finkel, 2024; Kellner, 2024). Thus, the online presence of the Otolith 351 Collection is a vital step toward creating a national platform that supports the analysis of 352 bony fish otoliths in various contexts, including paleontological, archaeological, biological, 353 and ecological studies (Begg et al., 2005; Disspain et al., 2016). 354 355

356 Otoliths serve multiple research purposes, including paleontological, archaeological, and 357 biological explorations, and support species identification in archaeological contexts. The 358 Otolith Collection of the Gulf of Mexico and Caribbean Sea represents the ichthyological 359 diversity of the region, encompassing 214 species of 67 families, reflecting over 11% of 360 the total species richness in the region. The collection's species composition mirrors the 361 local ichthyofauna's ecological importance and its significance to the regional economy, 362 making it a crucial resource for biodiversity studies. 363 364 With specimens collected from various regions, the collection provides valuable data for 365 studying species distribution, population variability, and temporal trends in the Gulf of 366 Mexico and the Caribbean. Furthermore, digitizing the Otolith Collection enhances global 367 accessibility, facilitates international collaboration, and supports research on critical global 368 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 challenges, such as climate change and conservation. The collection's website offers 369 innovative image processing software that improves species identification and taxonomic 370 studies, making these tools widely accessible to researchers. It is important to note that 371 while digitization expands access to the collection, physical specimens remain the primary 372 source of verifiable data, highlighting the need for their preservation. Therefore, we 373 emphasize the importance of adequate funding for maintaining this and other biological 374 collections, as they contribute significantly to knowledge, education, and innovation. 375 376 Acknowledgments 377 The authors wish to express their gratitude to the many students (especially Kylie López 378 Castillejos, Alt Gerardo Palacios Díaz, Diego Muñoz Pérez, José Emilio García Pérez, 379 Jonathan de la Cruz), fishermen (especially Fernando Mex and Enrique Mex), and 380 colleagues (especially Daniel Arceo Carranza, Claudia Durruty Lagunes, Manuel 381 Valenzuela Jiménez) who participated in the collection of scientific material, laboratory 382 work, analysis, and curatorship of the Otolith Collection from the Gulf of Mexico and 383 Caribbean Sea. We appreciate the generous donation of the Mexican Atlantic Fish Otolith 384 Collection by José Antonio Martínez Pérez from the Facultad de Estudios Superiores 385 Iztacala, Universidad Nacional Autónoma de México. This project was funded by PAPIIT-386 DGAPA-UNAM (IG201121). 387 388

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DOI:  https://doi.org/10.3897/arphapreprints.e134421 National Academies of Sciences, Engineering, and Medicine (2020). Biological 504 Collections: Ensuring Critical Research and Education for the 21st Century. The National 505 Academies Press, Washington, D.C., 229 p. https://doi.org/10.17226/25592 506 507 Nolf D (1985) Otolithi piscium. In H. Schultze (Ed.) Handbook of Palaeoichthyology, 508 Volume 10. Gustav Fisher Verlag, New York, 145 p. 509 510 Oré-Villalba DO (2017). Catálogo fotográfico de otolitos de peces marinos y 511 dulceacuícolas del Perú. Boletín del Instituto del Mar de Perú, 32(2): 136-213. 512 https://repositorio.imarpe.gob.pe/bitstream/20.500.12958/3190/1/Boletin%2032%282%29-513 1.pdf 514 515 Platt C, Popper AN (1981) Fine Structure and Function of the Ear. In: Tavolga WN, 516 Popper AN, Fay RR (Eds.) Hearing and Sound Communication in Fishes. Proceedings in 517 Life Sciences. Springer, New York, NY. https://doi.org/10.1007/978-1-4615-7186-5_1. 518 519 Rivaton J, Bourret P (1999) Les otolithes des poisons de l'Indo-Pacifique. Documents 520 Scientifiques et Techniques II 2. Institut de recherche pour le développement, Nouvelle-521 Caledonie, 378 p. https://horizon.documentation.ird.fr/exl-doc/pleins_textes/divers09-522 06/010021515.pdf 523 524 Robertson DR, Van Tassell J (2023) Shorefishes of the Greater Caribbean: online 525 information system. Version 3.0 Smithsonian Tropical Research Institute, Balboa, 526 Panamá. https://biogeodb.stri.si.edu/caribbean/en/pages 527 528 Rojo AL (2015) Report in the Fish Otolith Collection at the Nova Scotia Museum. 529 Curatorial Report Number 105, Nova Scotia Museum, Halifax, 176 p. ISBN 978-1-55457-530 676-0 https://ojs.library.dal.ca/NSM/article/view/6440/5676 531 532 Soltis, P. S. 2017. Digitization of herbaria enables novel research. American Journal of 533 Botany 104(9):1281-1284. https://doi.org/10.3732/ajb.1700281 534 535 Suarez AV, Tsutsui ND (2004) The value of museum collections for research and society. 536 BioScience 54(1): 66-74. https://doi.org/10.1641/0006-537 3568(2004)054[0066:TVOMCF]2.0.CO;2 538 539 Tuset VM, Lombarte A, Assis CA (2008) Otolith atlas for the western Mediterranean, north 540 and central eastern Atlantic. Scientia Marina 72: 7-198 541 https://doi.org/10.3989/scimar.2008.72s17 542 543 Van der Laan R, Fricke R, Eschmeyer WN (eds) (2024) Eschmeyer's Catalog of Fishes: 544 Classification (http://www.calacademy.org/scientists/catalog-of-fishes-classification). 545 Electronic version accessed 21/07/2024. 546 547 Vollmar A, Macklin JA, Ford L (2010) Natural history specimen digitization: challenges 548 and concerns. Biodiversity informatics 7(2):93-112 https://doi.org/10.17161/bi.v7i2.3992 549 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 550 Volpedo AV, Echeverría D (2000) Catálogo y claves de otolitos para la identificación de 551 especies del mar argentino. Peces de importancia económica. Editorial Dunken, 552 Argentina, 90 p. ISBN: 987-518-353-9 553 554 Volpedo AV, Echeverría D (2003) Ecomorphological patterns of the sagitta in fish on the 555 continental shelf off Argentine. Fisheries Research 60(2-3): 551-560 556 https://doi.org/10.1016/S0165-7836(02)00170-4 557 558 559 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Figure 560 561 562 Figure 1. Diversity of sagittae otoliths from the Collection of Otoliths of Fishes from the 563 Gulf of Mexico and the Caribbean Sea: 1) Family Elopidae, Elops saurus Linnaeus, 564 1766, 2) Family Megalopidae, Megalops atlanticus (Valenciennes, 1847) 3) Family 565 Lepisosteidae, Atractosteus tropicus (Gill, 1863), 4) Family Albulidae, Albula vulpes 566 (Linnaeus, 1758), 5) Family Engraulidae, Anchoa cayorum (Fowler, 1906), 6) Family 567 Clupeidae, Dorosoma petenense (Günther, 1867), 7) Family Clupeidae, Cetengraulis 568 edentulus (Cuvier, 1829), 8) Family Characidae, Astyanax altior Hubbs, 1936, 9) Family 569 Heptapteridae, Rhamdia guatemalensis (Günther, 1864), 10) Family Ariidae, Bagre 570 marinus (Mitchill, 1815), 11) Family Synodontidae, Saurida brasiliensis Norman, 1935, 571 12) Family Phycidae, Urophycis floridana (Bean & Dresel, 1884), 13) Family 572 Holocentridae, Holocentrus adscensionis (Osbeck, 1765), 14) Family Ophidiidae, 573 Brotula barbata (Bloch & Schneider, 1801), 15) Family Batrachoididae, Sanopus 574 reticulatus Collette, 1983, 16) Family Pomatomidae, Pomatomus saltatrix (Linnaeus, 575 1766), 17) Family Scombridae, Scomberomorus maculatus (Mitchill, 1815) (Bloch, 576 1792), 22) Family Gobiidae, Awaous banana (Valenciennes, 1837), 23) Family 577 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Centropomidae, Centropomus parallelus Poey, 1860, 24) Family Sphyraenidae, 578 Sphyraena barracuda (Edwards, 1771), 25) Family Polynemidae, Polydactylus 579 octonemus (Girard, 1858), 2 6) Family Cyclopsettidae, Cyclopsetta chittendeni Bean, 580 1895, 27) Family Paralichthyidae, Citharichthys spilopterus Gunther, 1862, 28) Family 581 Achiridae, Achirus lineatus (Linnaeus, 1758), 29) Family Carangidae, Hemicaranx 582 amblyrhynchus (Cuvier, 1833), 30) Family Echeneidae, Echeneis neucratoides (Zuiew, 583 1786). 584 585 586 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 587 Figure 1. ...continuation 1) Family Rachycentridae, Rachycentron canadum (Linnaeus, 588 1766), 2) Family Cichlidae, Mayaheros urophthalmus (Gunther, 1862), 3) Family 589 Pomacentridae, Abudefduf saxatilis (Linnaeus, 1758), 4) Family Atherinopsidae, 590 Menidia colei Hubbs, 1936, 5) Family Rivulidae, Cynodonichthys tenuis (Meek, 1904), 591 6) Family Fundulidae, Fundulus grandissimus Hubbs, 1936), 7) Family Cyprinodontidae, 592 Cyprinodon artifrons Hubbs, 1936, 8) Family Poeciliidae, Belonesox belizanus Kner, 593 1860, 9) Family Belonidae, Strongylura timucu (Walbaum, 1792), 10) Family 594 Hemiramphidae, Hemirramphus brasiliensis (Linnaeus, 1758), 11) Family Mugilidae, 595 Mugil cephalus Linnaeus, 1758, 12) Family Serranidae, Diplectrum formosum 596 (Linnaeus, 1766), 13) Family Epinephelidae, Epinephelus adscensionis (Osbeck, 1765), 597 14) Family Grammistidae, Rypticus maculatus Holbrook, 1855, 15) Family Labridae, 598 Lachnolaimus maximus Walbaum, 1792, 16) Family Triglidae, Prionotus punctatus 599 (Bloch, 1793 ), 17) Family Scorpaenidae, Scorpaena plumieri Bloch, 1789, 18) Family 600 Kyphosidae, Kyphosus sectatrix (Linnaeus, 1766), 19) Family Priacanthidae, 601 Priacanthus arenatus Cuvier, 1829, 20) Family Malacanthidae, Caulolatilus cyanops 602 Poey, 1866, 21) Family Lutjanidae, Lutjanus griseus (Linnaeus, 1758), 22) Family 603 Gerreidae, Gerres cinereus (Walbaum, 1792), 23) Family Haemulidae, Haemulon 604 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 plumierii (Lacepède, 1801), 24) Family Sparidae, Lagodon rhomboides (Linnaeus, 605 1766), 25) Family Sciaenidae, Cynoscion nebulosus (Cuvier, 1830), 26) Family 606 Pomacanthidae, Pomacanthus arcuatus (Linnaeus, 1758), 27) Family Chaetodontidae, 607 Chaetodon ocellatus Bloch, 1781, 28) Family Acanthuridae, Achirus lineatus (Linnaeus, 608 1758), Acanthurus coeruleus Bloch & Schneider, 1801, 29) Family Tetraodontidae, 609 Sphoeroides testudineus (Linnaeus, 1758), 30) Family Balistidae, Balistes capriscus 610 Gmelin, 1789. 611 612 613 Table 614 615 Table 1. List of species of the Otolith Collection of the Gulf of Mexico and the Caribbean 616 Sea. Taxonomic classification according to Van der Laan et al. (2024). The size range 617 of the specimens corresponds to the total length (TL), except in some cases where the 618 standard length (SL) is used (*). 619 620 Family Species Number of specimens Size range (mm) Lepisosteidae Atractosteus tropicus (Gill, 1863) 2 540-593 Elopidae Elops saurus Linnaeus, 1766 38 87-613 Megalopidae Megalops atlanticus (Valenciennes, 1847) 2 *385-426 Albulidae Albula vulpes (Linnaeus, 1758) 6 351-292 Ophichthidae Ahlia egmontis (Jordan, 1884) 1 365 Engraulidae Anchoa cayorum (Fowler, 1906) 7 90 Anchoa cubana (Poey, 1868) 2 *55 Anchoa lamprotaenia (Hildebrand, 1943) 34 89-130 Cetengraulis edentulus Cuvier, 1829 9 *94-140 Clupeidae Dorosoma petenense (Günther, 1867) 6 *104-140 Harengula clupeola (Cuvier, 1829) 10 51.47-130 Harengula jaguana Poey, 1865 15 110-160 Opisthonema oglinum (Lesuer, 1818) 24 33-250 Sardinella aurita Valenciennes, 1847 3 *156-163 Cyprinidae Ctenopharyngodon idella (Valenciennes, 1844) 1 261 Characidae Astyanax altior Hubbs, 1936 23 55.7-79.1 Heptapteridae Rhamdia guatemalensis (Günther, 1864) 4 91.5-1293 Ariidae Ariopsis felis (Linnaeus, 1766) 108 56-402 Bagre marinus (Mitchill, 1815) 106 267-540 Synodontidae Saurida brasiliensis Norman, 1935 2 *184-203 Synodus foetens (Linnaeus, 1766) 22 52.4-376 Phycidae Urophycis floridana (Bean & Dresel, 1884) 5 256-327 Holocentridae Holocentrus adscensionis (Osbeck, 1765) 5 92-280 Holocentrus rufus (Walbaum, 1792) 2 110-115 Myripristis jacobus Cuvier,1829 2 166-180 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Ophidiidae Brotula barbata (Bloch & Schneider, 1801) 3 372-550.8 Ophidion holbrookii Putnam, 1874 3 77-141 Batrachoididae Opsanus beta (Goode & Bean, 1880) 6 *98-205 Opsanus dichrostomus Collette, 2001 2 105-130 Sanopus reticulatus Collette, 1983 12 280-590 Pomatomidae Pomatomus saltatrix (Linnaeus, 1766) 8 420-524 Scombridae Euthynnus alletteratus (Rafinesque, 1810) 1 450 Scomberomorus brasiliensis Collette, Russo & Zavala-Camin, 1978 5 *290-360 Scomberomorus cavalla (Cuvier, 1829) 4 695-750 Scomberomorus maculatus (Mitchill, 1815) 4 480-590 Trichiuridae Trichiurus lepturus (Linnaeus, 1758) 8 *268-884 Dactylopteridae Dactylopterus volitans (Linnaeus, 1758) 3 86-92 Mullidae Mulloidichthys martinicus (Cuvier,1829) 2 250-268 Pseudopenaeus maculatus (Bloch, 1793) 1 223 Upeneus parvus Poey, 1852 3 *115-145 Callionymidae Chalinops pauciradiatus (Gill, 1865) 6 27.2-36.9 Eleotridae Dormitator maculatus (Bloch, 1792) 6 45-90 Gobiomorus dormitor Lacepéde, 1800 6 *147-186 Gobiidae Awaous banana (Valenciennes, 1837) 2 *51-62 Gobionellus oceanicus (Pallas, 1770) 5 *93-159 Gobiosoma robustum (Ginsburg, 1933) 6 28.2-36.3 Centropomidae Centropomus parallelus Poey, 1860 3 *158-181 Centropomus undecimalis (Bloch, 1792) 25 370-473 Sphyraenidae Sphyraena barracuda (Edwards, 1771) 22 247-1015 Sphyraena guachancho Cuvier, 1829 9 *148-583 Polynemidae Polydactylus octonemus (Girard, 1858) 3 *78-195 Cyclopsettidae Cyclopsetta chittendeni Bean, 1895 117 218-315 Cyclopsetta fimbriata (Goode & Bean, 1885) 51 77-390 Syacium papilosum (Linnaeus, 1758) 34 231-275 Paralichthyidae Citharichthys macrops Dressel, 1885 1 153 Citharichthys spilopterus Gunther, 1862 5 *56-138 Paralichthys albigutta (Jordan & Gilbert, 1882) 2 139-530 Achiridae Achirus lineatus (Linnaeus, 1758) 10 60-93.3 Trinectes maculatus (Bloch & Schneider, 1801) 1 94.4 Carangidae Alectis ciliaris (Bloch, 1787) 1 302 Caranx crysos (Mitchill, 1815) 27 271-405 Caranx hippos (Linnaeus, 1766) 17 233-370 Caranx latus Agassiz, 1831 8 121-259 Caranx ruber (Bloch, 1793) 4 246-305 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Chloroscombrus chrysurus (Linnaeus, 1766) 6 *17-182 Hemicaranx amblyrhynchus (Cuvier, 1833) 1 *240 Oligoplites saurus Bloch & Schneider, 1801 16 106-278 Selar crumenophthalmus (Bloch, 1793) 6 *185-207 Selene setapinnis (Mitchill, 1815) 5 *149-185 Selene vomer (Linnaeus, 1758) 7 305-315 Trachinotus carolinus (Linnaeus, 1766) 3 300-305 Trachinotus falcatus (Linnaeus, 1758) 21 96-520 Trachinotus goodei (Jordan & Evermann, 1896) 5 218-295 Trachinotus meeki Brind, 1918 1 116.1 Echeneidae Echeneis neucratoides (Zuiew, 1786) 10 320-523 Rachycentridae Rachycentron canadum (Linnaeus, 1766) 3 219-690 Cichlidae Mayaheros urophthalmus (Gunther, 1862) 11 123.9-235 Oreochromis niloticus (Linnaeus, 1758) 1 245 Parachromis friedrichsthalii (Heckel, 1840) 3 103-113 Parachromis motaguensis (Günter, 1867) 4 230-269 Petenia splendida (Günter, 1862) 5 238-265 Rocio octofasciata Regan, 1903 10 48-74.5 Thorichthys meeki Brind, 1918 3 54.1-96.7 Pomacentridae Abudefduf saxatilis (Linnaeus, 1758) 5 75.4-156 Neopomacentrus cyanomos (Bleeker, 1856) 10 32.1-85.8 Stegastes leucostictus (Müller & Troschel, 1848) 2 106.5- 109.5 Stegastes xanthurus (Poey, 1860) 2 94.6-97.7 Atherinopsidae Atherinomorus stipes (Muller & Troschel, 1848) 21 38.4-56.63 Menidia colei Hubbs, 1936 9 11.5-33 Menidia sp. 12 19.6-24.38 Rivulidae Cynodonichthys tenuis (Meek, 1904) 1 41 Kryptolebias marmoratus (Poey, 1880) 1 30 Fundulidae Fundulus grandissimus Hubbs, 1936 41 90.6-160 Fundulus persimilis Miller, 1955 19 78.6-122.2 Lucania parva (Baird & Girard, 1855) 9 24-35.3 Cyprinodontidae Cyprinodon artifrons Hubbs, 1936 57 16.4-45 Floridichthys polyommus Hubbs, 1936 43 62-120 Jordanella pulchra Hubbs, 1936 14 28-35.4 Poeciliidae Belonesox belizanus Kner, 1860 12 55.9-91.5 Gambusia yucatana Regan, 1914 14 19.8-36.5 Poecilia mexicana (Steindachner, 1863) 3 74.1-88.8 Poecilia velifera (Regan, 1914) 18 38-79 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Pseudoxiphophorus bimaculatus (Heckel 1848) 2 55-59.8 Belonidae Strongylura marina (Walbaum, 1792) 10 260-332 Strongylura notata (Poey, 1860) 42 195-433 Strongylura timucu (Walbaum, 1792) 9 230 Tylosurus crocodilus (Peron y Lesueur, 1821) 11 65-1020 Hemiramphidae Chriodorus atherinoides Goode & Bean, 1882 28 109-190 Hemirramphus brasiliensis (Linnaeus, 1758) 6 300 Hyporhamphus unifasciatus (Ranzani, 1841) 43 140-326 Mugilidae Mugil cephalus Linnaeus, 1758 14 127-232 Mugil curema Valenciennes,1836 17 120-434 Mugil trichodon Poey,1875 8 39.7-183 Serranidae Diplectrum formosum (Linnaeus, 1766) 26 125-248 Hypoplectrus ecosur Victor, 2012 7 58.1-115 Serranus subligarius (Cope, 1870) 7 64.2-92 Epinephelidae Cephalopholis cruentata (Lacepede, 1802) 6 270-305 Cephalopholis fulva (Linnaeus, 1758) 1 235 Epinephelus adscensionis (Osbeck, 1765) 12 *31-256 Epinephelus morio (Valenciennes, 1828) 14 270-480 Hyporthodus niveatus (Valenciennes, 1828) 3 350-367 Mycteroperca bonaci (Poey, 1860) 3 345-625 Grammistidae Rypticus maculatus Holbrook, 1855 3 140-142 Labridae Halichoeres radiatus (Linnaeus, 1758) 1 *264 Lachnolaimus maximus Walbaum, 1792 23 140-325 Nicholsina usta (Valenciennes, 1840) 4 95-165 Scarus coeruleus (Edwards, 1771) 1 130 Sparisoma rubripinne (Valenciennes, 1840) 2 385 Triglidae Prionotus punctatus (Bloch, 1793) 2 280- Prionotus scitulus Jordan & Gilbert, 1882 2 214-215 Prionotus tribulus Cuvier, 1829 4 175-300 Scorpaenidae Pterois volitans (Bloch, 1758) 1 *95.2 Scorpaena brasiliensis Cuvier, 1829 5 155-215 Scorpaena plumieri Bloch, 1789 14 105-240 Kyphosidae Kyphosus sectatrix (Linnaeus, 1766) 4 140-427 Priacanthidae Priacanthus arenatus Cuvier, 1829 3 310-325 Pristigerys alta (Gill, 1862) 1 45 Malacanthidae Caulolatilus cyanops Poey, 1866 1 *290 Lutjanidae Lutjanus analis (Cuvier, 1828) 2 398 Lutjanus apodus (Walbaum,1792) 1 206 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Lutjanus buccanella (Cuvier, 1828) 2 *180-191 Lutjanus campechanus (Poey, 1860) 3 455-470 Lutjanus cyanopterus (Cuvier,1828) 1 206 Lutjanus griseus (Linnaeus, 1758) 20 69.2 Lutjanus jocu (Bloch & Schneider, 1801) 1 348 Lutjanus synagris (Linnaeus, 1758) 38 185-405 Ocyurus chrysurus (Bloch, 1791) 24 68.9-321 Rhomboplites aurorubens (Cuvier, 1829) 15 260-277 Gerreidae Diapterus auratus Ranzani, 1842 6 *134-173 Diapterus rhombeus (Cuvier, 1829) 5 *116-139 Eucinostomus argenteus (Baird & Girard, 1855) 4 96.4-117.5 Eucinostomus gula (Quoy & Gaimard, 1824) 42 66-113 Eucinostomus harengulus Goode & Bean, 1880 5 *61-95 Eucinostomus melanopterus (Bleeker, 1863) 2 *61-56 Eugerres brasilianus (Cuvier, 1830) 7 180-280 Eugerres plumieri (Cuvier, 1830) 45 96-317 Gerres cinereus (Walbaum, 1792) 26 107.4-265 Haemulidae Anisotremus virginicus (Linnaeus, 1758) 25 181-367 Conodon nobilis (Linnaeus, 1758) 6 *57-185 Haemulon aurolineatum Cuvier, 1830 24 162-249 Haemulon bonariense Cuvier, 1830 8 150-342 Haemulon carbonarium Poey, 1860 6 *147-265 Haemulon flavolineatum (Desmarest, 1823) 6 *155-185 Haemulon parra (Desmarest, 1823) 3 *215-259 Haemulon plumierii (Lacepède, 1801) 102 160-315 Haemulon sciurus (Shaw,1803) 4 186-253 Orthopristis chrysoptera (Linnaeus, 1766) 26 127.1-244 Rhonciscus crocro (Cuvier, 1830) 2 *192-211 Sparidae Archosargus probatocephalus (Walbaum, 1792) 24 176-610 Archosargus rhomboidalis (Linnaeus, 1758) 67 85.1-280 Calamus bajonado (Bloch & Schneider, 1801) 15 215-540 Calamus calamus (Valenciennes, 1830) 11 191-300 Calamus campechanus Randall & Caldwell, 1966 18 185-249 Calamus nodosus Randall & Caldwell, 1966 6 229-310 Calamus pennatula Guichenot, 1868 3 282 Calamus proridens Jordan & Gilbert, 1884 5 210-275 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Lagodon rhomboides (Linnaeus, 1766) 31 44.5-207 Stenotomus chrysops (Linnaeus, 1766) 1 *210-521 Sciaenidae Bairdiella chrysoura (Lacepéde, 1802) 5 82.2-165 Bairdiella ronchus (Cuvier, 1830) 19 139-270 Corvula batabana (Poey, 1860) 2 156.6-160 Cynoscion arenarius Ginsburg, 1930 92 160-423 Cynoscion jamaicensis (Vaillant & Bocourt, 1883) 2 *256-268 Cynoscion nebulosus (Cuvier, 1830) 92 110-490 Cynoscion nothus (Holbrook, 1848) 6 130-245 Eques lanceolatus Linnaeus, 1758 32 80-335 Menticirrhus americanus (Linnaeus, 1758) 191 204-400 Menticirrhus littoralis (Holbrook, 1847) 11 91-365 Menticirrhus saxatilis (Bloch &Schneider, 1801) 3 *226-236 Micropogonias furnieri (Desmarest,1823) 59 74-412 Micropogonias undulatus (Linnaeus, 1766) 39 86-425 Pareques umbrosus Jordan & Eigenmann, 1889 7 55-210 Pogonias cromis (Linnaeus, 1766) 1 84 Stellifer lanceolatus (Holbrook, 1855) 6 *75-96 Umbrina coroides Cuvier, 1830 7 *80-214 Lobotidae Lobotes surinamensis (Bloch, 1790) 6 320-570 Pomacanthidae Holacanthus bermudensis Goode, 1876 8 65-337 Pomacanthus arcuatus (Linnaeus, 1758) 5 87-498 Chaetodontidae Chaetodon ocellatus Bloch, 1781 34 40-124 Ephippidae Chaetodipterus faber (Broussonet, 1782) 16 63.8-354 Acanthuridae Acanthurus chirurgus (Bloch, 1787) 4 316 Acanthurus coeruleus Bloch & Schneider, 1801 5 201-255 Acanthurus tractus Poey, 1860 5 244-325 Ogcocephalidae Ogcocephalus cubifrons (Richardson, 1836) 2 125-215 Diodontidae Chilomycterus schoepfii (Walbaum, 1792) 13 96-285 Diodon hystrix (Linnaeus, 1758) 1 490 Tetraodontidae Lagocephalus laevigatus (Linnaeus, 1766) 5 330-554 Sphoeroides nephelus (Goode & Bean, 1882) 6 124.6-280 Sphoeroides spengleri (Bloch, 1785) 8 54-111.2 Sphoeroides testudineus (Linnaeus, 1758) 7 157-235 Ostraciidae Acanthostracion quadricornis (Linnaeus, 1758) 11 90-270 Monacanthidae Monacanthus ciliatus (Mitchill, 1818) 4 81-136 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421 Stephanolepsis hispida (Linnaeus, 1766) 2 160 Balistidae Balistes capriscus Gmelin, 1789 10 240-495 621 622 Author contribution 623 Maribel Badillo Alemán: Conceptualization, investigation, data curation, supervision, 624 writing original draft. 625 Ariana Solís Gómez: Investigation, data curation. 626 Alfredo Gallardo Torres: Investigation, data curation, formal analysis, resources. 627 Eduardo Pacheco Gongora: Data curation, software development, visualization. 628 Xavier Chiappa-Carrara: Conceptualization, investigation, project administration, formal 629 analysis, writing original draft, funding acquisition. 630 631 Author-formatted, not peer-reviewed document posted on 14/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e134421