Two new species of Hyalonema (Hexactinellida, Amphidiscosida, Hyalonematidae) from the Indo-West Pacific

81 Two new species of Hyalonema (Hexactinellida, Amphidiscosida, Hyalonematidae) from the Indo-West Pacific Lin Gong1,2 , Edwin Setiawan3, Xinzheng Li1,2,4 , Swee Cheng Lim5 1 Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China 2 College of Marine Science, University of Chinese Academy of Sciences, Beijing, China 3 Laboratory of Animal Bioscience and Technology, Department of Biology, Institut Teknologi Sepuluh Nopember, Surabaya, Indonesia 4 Laboratory for Marine Biology and Biotechnology, Qingdao National Laboratory for Marine Science and Technology, Qingdao, China 5 Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, Singapore 117377, Singapore Corresponding author: Swee Cheng Lim ([email protected]) Copyright: © Lin Gong et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract This study describes two new species collected from deep-sea environments in the Indo-West Pacific. Hyalonema (Cyliconema) dexiangi sp. nov., collected from the Indian Ocean, is characterized by distinctive finger-like protuberances on the body surface, the presence of choanosomal pentactins, the co-occurrence of micropentactins and microhexactins, and the absence of mesamphidiscs. Hyalonema (Cyliconema) subconica sp. nov., sampled from a seamount in the western Pacific Ocean, is distinguished by the presence of pinular diactins, a unique combination of choanosomal spicules (diactins, pentactins, and hexactins), and the presence of basalia bearing two-toothed anchors. In addition to their morphological distinctions, molecular phylogenetic analyses based on 16S rDNA sequences indicate a close genetic relationship between the two species. Key words: Deep sea, glass sponge, Hyalonema (Cyliconema) dexiangi sp. nov., Hyalonema (Cyliconema) subconica sp. nov., new species, phylogeny, taxonomy Introduction Hyalonema Gray, 1832 is the most speciose genus within the class Hexactinellida, comprising 13 subgenera and 119 species (De Voogd et al. 2025). More than 90 species were described prior to 1930, based primarily on material collected during historical expeditions. These include the HMS Challenger global survey (1873–1876) (Schulze 1886, 1887), the Valdivia Expedition (1898–1899) off West Sumatra (Schulze 1904), and the Siboga Expedition (1899–1900) in the Banda Sea region (Ijima 1927). Distinct morphological characteristics differentiate some subgenera of Hyalonema (e.g., H. (Prionema) Lendenfeld, 1915; H. (Paradisconema) Ijima, 1927; and H. (Oonema) Lendenfeld, 1915), while diagnostic features remain ambiguous in certain subgenera, notably H. (Coscinonema) Ijima, 1927. Taxonomic revision of Hyalonema presents significant challenges due to its remarkable species richness and the historical status of many type specimens. Molecular studies on Hyalonema remain limited. To date, only 16S rDNA sequences from representatives of five subgenera Academic editor: Martin Dohrmann Received: 11 July 2025 Accepted: 6 November 2025 Published: 3 December 2025 ZooBank: https://zoobank. org/493A2920-0CAE-46BD-B96C91E4F2C7AFE5 Citation: Gong L, Setiawan E, Li X, Lim SC (2025) Two new species of Hyalonema (Hexactinellida, Amphidiscosida, Hyalonematidae) from the Indo-West Pacific. ZooKeys 1262: 81–96. https://doi.org/10.3897/ zookeys.1262.164821 ZooKeys 1262: 81–96 (2025) DOI: 10.3897/zookeys.1262.164821 82 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific (H. (Corynonema), H. (Prionema), H. (Cyliconemaoida), H. (Cyliconema), H. (Onconema)) have been analysed, with few molecular data available for species-level identification (Kersken et al. 2018a; Dohrmann 2019). Hyalonema (Cyliconema) Ijima, 1927 is characterized by dermal spicules bearing whip-like pinular rays, macramphidiscs with umbels broader than long, and the absence of both ambuncinates and a sieve-plate (Tabachnick and Menshenina 2002). The subgenus comprises 30 species with a broad geographical distribution (Fig. 1), based on the locations of their holotypes, spanning the Pacific, Atlantic, and Indian Oceans (Suppl. material 1). Notably, most species occur in the Indian Ocean, with over half recorded at depths exceeding 1000 m. The World Porifera Database (https://www.marinespecies.org/porifera/) lists relatively few species from the western Pacific Ocean. Six valid H. (Cyliconema) species were recorded from Indonesian waters: H. (Cyliconema) apertum Schulze, 1886 from the Banda Sea and West Sumatra; H. (Cyliconema) keiense Ijima, 1927, and H. (Cyliconema) timorense Ijima, 1927 collected during the Siboga Expedition in the Banda Sea; H. (Cyliconema) martabanense Schulze, 1900, H. (Cyliconema) rapa Schulze, 1900, and H. (Cyliconema) tulipa Schulze, 1904, all obtained during the Valdivia Expedition off West Sumatra. Species of Hyalonema predominantly inhabit muddy benthic environments across the world’s oceans (Tabachnick and Menshenina 2024) and have been reported in polymetallic nodule fields (Kersken et al. 2018b; Stratmann et al. 2021). In 2013, a research cruise conducted by the Institute of Oceanology, Figure 1. Type locations of species of Hyalonema (Cyliconema): 1. H. (Cyliconema) abyssale; 2. H. (Cyliconema) apertum; 3. H. (Cyliconema) clavapinulatum; 4. H. (Cyliconema) coniforme; 5. H. (Cyliconema) conqueror; 6. H. (Cyliconema) curvisclera; 7. H. (Cyliconema) drygalskii; 8. H. (Cyliconema) eupinnulum; 9. H. (Cyliconema) globiferum; 10. H. (Cyliconema) hozawai; 11. H. (Cyliconema) infundibulum; 12. H. (Cyliconema) keiense; 13. H. (Cyliconema) lanceolatum; 14. H. (Cyliconema) madagascarense; 15. H. (Cyliconema) martabanense; 16. H. (Cyliconema) masoni; 17. H. (Cyliconema) molle; 18. H. (Cyliconema) nicobaricum; 19. H. (Cyliconema) ovatum; 20. H. (Cyliconema) pirum; 21. H. (Cyliconema) polycaulum; 22. H. (Cyliconema) rapa; 23. H. (Cyliconema) simile; 24. H. (Cyliconema) somalicum; 25. H. (Cyliconema) tasmani; 26. H. (Cyliconema) tenerum; 27. H. (Cyliconema) thomsonis; 28. H. (Cyliconema) timorense; 29. H. (Cyliconema) tulipa; 30. H. (Cyliconema) valdiviae; 31. H. (Cyliconema) dexiangi sp. nov.; 32. H. (Cyliconema) subconica sp. nov. 83 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Chinese Academy of Sciences, to the Yap Seamount in the northwestern Pacific Ocean collected a single specimen of Hyalonema (Cyliconema) using the remotely operated vehicle (ROV) Fa Xian. A second specimen was obtained during the South Java Sea Expedition (SJADES) in 2018, jointly organized by the Lee Kong Chian Natural History Museum (LKCNHM) at the National University of Singapore (NUS) and the Indonesian Research Agency (BRIN). Integrated morphological and molecular analyses confirm that these specimens represent two new species, which are formally described and illustrated here. We also provide the corresponding 16S rDNA and COI sequences. However, given the current scarcity of molecular data for Hyalonema, establishing a comprehensive phylogenetic framework for the genus remains difficult. Material and methods One sponge sample was collected in the Indian Ocean using a beam trawl during the SJADES expedition from 23 March to 5 April 2018 (Chim et al. 2021; Ng and Rahayu 2021). The sample was deposited at the Lee Kong Chian Natural History Museum (LKCNHM), Singapore. One sponge sample was collected by the submersible ROV Fa Xian during a cruise of the research ship Ke Xue in the western Pacific Ocean. The sample was deposited in the Marine Biological Museum of the Chinese Academy of Sciences (MBM) at the Institute of Oceanology of the Chinese Academy of Sciences (IOCAS). Total genomic DNA was extracted using the Tissue DNA Kit (OMEGA Bio-Tek) according to manufacturer’s protocols. The libraries were finally sequenced using a next-generation sequencing platform (Illumina) with PE150 mode according to the standard protocols. Raw sequence reads were quality-trimmed using Trimmomatic (Bolger et al. 2014) to remove adaptor contaminants and low-quality reads. The clean reads were then assembled into a set of contigs for each sample using MegaHit v. 1.2.9 (Li et al. 2015). Finally, the 16S rDNA and COI genes were recovered from the assembled contigs. A phylogenetic tree was constructed from partial 16S rDNA sequences of Hyalonema. The reference sequence alignment for the 16S rDNA gene was obtained from the GitHub repository by Dohrmann et al. (2023) (https://github.com/ PalMuc/SONNE_Hexactinellida/blob/main/16S.fasta). This dataset was subsequently curated to exclude all representatives of the subclass Hexasterophora, with the pheronematid Semperella jialongae retained as the outgroup for rooting the tree. The untrimmed and trimmed alignments are provided in Suppl. materials 2, 3. We used the workflow desktop platform PhyloSuite (Zhang et al. 2020) to build the phylogenetic trees. For Bayesian inference (BI) and maximum-likelihood (ML) analyses, the best-fit substitution models were HKY+G and TIM2+R inferred by ModelFinder (Kalyaanamoorthy et al. 2017). For the ML analysis, we employed IQ-TREE v. 1.6.8 (Nguyen et al. 2015), with node support assessed through ultrafast bootstrap approximation (UFBoot) based on 1000 replicates. The BI tree was reconstructed using MrBayes v. 3.2 (Ronquist et al. 2012). The analysis consisted of two independent runs of four Markov chains each, each running for 10,000,000 iterations, with sampling every 1000 iterations. After discarding the first 25% of trees as burn-in, the remaining trees were used to construct the 50% majority-rule consensus tree and to estimate posterior probabilities (PPs). Phylogenetic tree annotation was performed using the webtool iTOL (https://itol.embl.de/). 84 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Results Taxonomy Family Hyalonematidae Gray, 1857 Genus Hyalonema Gray, 1832 Subgenus Hyalonema (Cyliconema) Ijima, 1927 Hyalonema (Cyliconema) dexiangi Gong, Setiawan & Lim, sp. nov. https://zoobank.org/0EAE2FA7-771B-4698-AEB2-14EA1ADBBAC8 Figs 2, 3, Table 1 Material examined. Holotype: ZRC.POR.0519. Station CP23 (6°46.739'S, 105°09.239'E). sample number CP2301. 30 March 2018, collected by Lim Swee-Cheng. Beam Trawl. Depth 571 m, gravel and mud substrates. Indian Ocean, south of Cilacap (West Java, Indonesia). Description. The sponge exhibits a cylindrical form (Fig. 2A, B) with finger-like protuberances on its surface (Fig. 2C), each terminating in an osculum (diameter 12 mm). The apex of the specimen exhibits a finger-like form without a terminal osculum or sieve-plate (Fig. 2E, F). Due to trawling collection, the specimen is morphologically damaged and lacks basalia (Fig. 2D). The specimen measures 224 mm in body length and 106 mm in width. The dermal surface displays relatively uniform mesh openings, ranging from 3 mm to 6 mm in diameter. Spicules. Atrialia are pinular pentactins (Fig. 3A) with longer spines on the pinular rays (170–282 μm in length) and tangential rays (17–41 μm in length) bearing small spines. Dermalia are pinular pentactins (Fig. 3B), elongated and whip-like, featuring pinular rays (117–384 μm in length) with longer spines and tangential rays (21–40 μm in length) bearing small spines. Canalaria are pinular pentactins (Fig. 3C) with spines on the pinular rays (96–367 μm in length), and tangential rays (23–52 μm in length) bearing small spines. The choanosomal skeleton consists of diactins and pentactins. The diactins (Fig. 3G) have smooth surfaces, pointed ends (Fig. 3F), and some exhibit a medial swelling (1140–2883 µm in length). Hypodermalia (Fig. 3H) and hypoatrialia are pentactins with smooth rays, and hypodermalia pentactins (257–442 μm in length) are more abundant than hypoatrialia pentactins (224–336 μm in length). Choanosomal pentactins (288–575 μm in length) with smooth rays are occasionally present, and hexactins are rarely observed. Pinular diactins, which are extremely scarce, have a pinular ray densely covered with spines (Fig. 3I), with the other end pointed and medially tuberculate, measuring 712–1029 μm in length. Microscleres are two types of amphidiscs, micropentactins and microhexactins. Macramphidiscs (Fig. 3J) have smooth disc surface and tuberculated axial surface, measuring 228–380 μm in length. The disc diameter (110–145 μm) is greater than the disc length (80–106 μm). Micramphidiscs (Fig. 3K) (14–21 μm in length) possess palmate heads and nearly smooth shafts sparsely covered with spines. Micropentactins (Fig. 3E) have four tangential rays (41–98 μm in length) of approximately equal length. Microhexactins (Fig. 3D) have rays (27– 86 μm in length) with slightly curved tips. Etymology. Dexiangi is named in honor of the late Professor Wang Dexiang from Xiamen University. A passionate and dedicated sponge researcher, Professor Wang made significant contributions to sponge science and mentored 85 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Figure 2. Hyalonema (Cyliconema) dexiangi sp. nov. A, B. External morphology of the specimen; C. Finger-like protuberances of the specimen; D. Holes left after the basalia torn off during bottom trawling; E, F. Atrial surface. 86 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Table 1. Measurements of the spicules of Hyalonema (Cyliconema) dexiangi sp. nov. (in µm); N = number of spicules measured; s.d. = standard deviation; range = range from the minimum to the maximum. Nmean range s.d. Dermalia, pinule pinular ray length 20 248 117–384 89 pinular ray width 20 8 5–11 2 tangential ray length 20 32 21–40 5 tangential ray width 20 64–10 1 Atrialia, pinule pinular ray length 20 215 170–282 32 pinular ray width 20 7 6–8 0.7 tangential ray length 20 27 17–41 6 tangential ray width 20 5 4–7 0.7 Canalaria, pinule pinular ray length 20 219 96–367 79 pinular ray width 20 74–10 2 tangential ray length 20 34 23–52 8 tangential ray width 20 6 3–7 1 Hypodermalia, pentactin tangential ray length 20 349 257–442 47 tangential ray width 20 18 14–21 2 Hypoatrialia, pentactin tangential ray length 8 294 224–336 35 tangential ray width 8 15 12–17 2 Choanosomalia, diactin length 20 1812 1140–2883 512 width 20 13 9–18 13 Choanosomalia, pentactin tangential ray length 15 384 276–600 93 tangential ray width 15 21 15–44 7 Pinular diactin length 3 824 712–1029 178 width 3 8 7–8 0.4 Macramphidisc length 12 329 228–380 362 umbel length 12 95 80–106 8 umbel diameter 12 131 110–145 10 Micramphidisc length 20 17 14–21 2 umbel length 20 6 4–8 1 umbel diameter 20 6 5–8 0.7 Microhexactin length 20 62 41–98 13 Micropentactin length 20 61 27–86 15 87 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Figure 3. SEM images of spicules of Hyalonema (Cyliconema) dexiangi sp. nov. A. Atrial pinular pentactin; B. Dermal pinular pentactin; C. Canalarial pinular pentactin; D. Microhexactin; E. Micropentactin; F. Details of diactin; G. Diactin; H. Dermal pentactin; I. Pinular diactin; J. Macramphidisc; K. Micramphidisc. numerous students in China. Remembered for his generosity, kindness, and collegial spirit, his passing represents a profound loss to the scientific community and all who knew him. Type locality. Indian Ocean south of Java, 571 m. GenBank number. PV625033 (16S rDNA) and PV618251 (COI). Molecular data. This study obtained molecular sequences of two new species of H. (Cyliconema). In the Bayesian inference (BI) tree based on 16S rDNA sequences (Fig. 4), the new species groups with another new species described below. Currently available Hyalonema sequences in GenBank represent five subgenera. Phylogenetic analysis revealed that H. (Cyliconema) groups with H. (Cyliconemaoida), along with five unidentified Hyalonema species and Lophophysema eversa Gong, Li & Qiu, 2014. The topology of the max- 88 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Figure 4. Phylogenetic tree obtained by Bayesian inference (BI) analysis based on 16S rDNA. The numbers at each node are Bayesian posterior probabilities (PP, left) and UFBoot (right) values. Support values under 50% are not shown. Semperella jialongae Hyalonema Onconema agassizi Hyalonema Corynonema depressum Hyalonema Hyalonema Cyliconemaoida ovuliferum Hyalonema Prionema breviradix Hyalonema 1.0/92.9 Hyalonema 1.0 Hyalonema Hyalonema Cyliconemaoida campanula Hyalonema (Cyliconema) subconica sp. nov. Hyalonema (Cyliconema) dexiangi sp. nov. 0.5/86.3 1.0 Lophophysema eversa Hyalonema Tabachnickia Hyalonema Onconema obtusum Hyalonema Onconema clarioni Tree scale: 0.1 Genus Semperella Hyalonema Corynonema Hyalonema Prionema Hyalonema Hyalonema Cyliconemaoida Hyalonema Onconema Tabachnickia Lophophysema Hyalonema Cyliconema) 0.6/79.1 0.5/79.1 1.0/91.6 0.8 0.6 0.8 1.0/95.8 • not supported by ML analysis • • • imum-likelihood (ML) tree was largely congruent with that of the BI tree, as shown in Suppl. material 4. Remarks. The new species exhibits dermal spicules with whip-like pinular rays, macramphidiscs displaying umbels broader than long, and an absence of ambuncinates and sieve-plate, which align with the diagnostic characteristics of Hyalonema (Cyliconema). Within the subgenus Hyalonema (Cyliconema), only three species – H. (Cyliconema) ovatum Ijima, 1895, H. (Cyliconema) tulipa Schulze, 1904 and H. (Cyliconema) valdiviae Schulze, 1904 – lack mesamphidiscs, a trait consistent with the new species. However, the new species possesses micropentactins, which have not been reported in these three species. Notably, H. (Cyliconema) tulipa, also recorded in Indonesian waters, exhibits strongly curved rays of microhexactins and features a vertically protruding central cone on its atrial surface. In contrast, the microhexactins of the new species show only slight curvature at the ray tips and lack a central cone on the atrial surface. Externally, the sponge exhibits finger-like protuberances on its surface, a morphology closely resembling that of Composocalyx Schulze, 1904, which distinguishes it from species within the genus Hyalonema. The new species is further characterized by the presence of choanosomal pentactins, a feature reported in only a few species, such as H. (Onconema) clarioni Kersken, Janussen & Martínez Arbizu, 2018 and H. (Prionema) breviradix Kersken, Janussen & Martínez Arbizu, 2018, and this distinguishes it readily from other Hyalonema species. These morphological differences confirm the status of this specimen as a new species. 89 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Hyalonema (Cyliconema) subconica Gong, Li & Lim, sp. nov. https://zoobank.org/165AFAC1-9F1A-46F3-B472-6B937C859E8D Figs 5–7, Table 2 Material examined. Holotype: MBM288250. 15 December 2014, Depth 1106 m, foraminiferal ooze substrate. Yap seamount (8°51.66'N, 137°44.09'E). Description. The sponge is white, anchored to the seafloor of foraminiferal ooze by a cluster of intertwined basalia (Fig. 5A, B). The sponge is bell-shaped, resembling a rose (Fig. 5C). The surface of the basalia is colonized by anemones (Fig. 5D). The basalia measure nearly 39 cm in length, while the main body of the sponge is 8 cm long, with an atrial surface diameter of 6 cm. The atrial surface has relatively uniform mesh openings (Fig. 6A), with pore diameters ranging from 0.3 mm to 2.3 mm. The mesh openings are separated by non-porous tissue of uneven distribution, varying in width from 0.5 mm to 5 mm. The dermal areas display distinct mesh patterns (Figs 5E, 6B) that are smaller than those of the sieve-plates, with pore diameters 0.1 mm to 0.6 mm. Spicules. The choanosomal skeleton primarily consists of diactins (Fig. 7B) (893–1977 µm in length), which have smooth surfaces, pointed ends, and occasionally exhibit medial swellings. Pentactins (ray length: 288–575 μm) and hexactins (ray length: 270–377 μm) with smooth rays are occasionally observed. Dermalia are pinular pentactins (Fig. 7D), elongated and whip-like, featuring pinular rays (206–277 μm in length) with short spines and tangential rays (18– 29 μm in length) bearing small spines. Atrialia are pinular pentactins (Fig. 7E) with longer spines on the pinular rays (147–252 μm in length) and tangential rays (23–40 μm in length) bearing small spines. Hypodermalia (288–575 μm in length) and hypoatrialia (266–624 μm in length) are pentactins with smooth rays, like choanosomal pentactins (Fig. 7A). Canalaria are pinular pentactins (Fig. 7F) with spines on the pinular rays (186–240 μm), and tangential rays (22–36 μm in length) bearing small spines. Pinular diactins (273–708 μm in length) have a pinular ray densely covered with spines (Fig. 7C), with the other end rounded or pointed, medially tuberculate. The basalia were predominantly broken during collection, making it impossible to observe the terminal structures. And, on rare occasions, one or two slender spicules were observed to display two-toothed anchors (Fig. 7K). Microscleres are three types of amphidiscs and micropentactins. Macramphidiscs (Fig. 7G) are oval-shaped, measuring 220–244 μm in length, with smooth disc surfaces and tuberculated axial surface. The disc diameter (80– 101 μm) is greater than the disc length (56–73 μm). Mesamphidiscs (Fig. 7H) measure 39–69 μm in length, with more teeth on the terminal discs compared to macramphidiscs. The axial surface exhibits numerous tubercles, and the disc diameter (11–19 μm) is smaller than the disc length (13–27 μm). Micramphidiscs (Fig. 7I) (13–21 μm in length) have disc diameters (4–(5)–7 μm) smaller than disc lengths (4–(6)–7 μm), and the teeth are densely arranged. Micropentactins (Fig. 7F), which are extremely scarce, possess four tangential rays (37–76 μm in length) of approximately equal length. We observed only one microhexactin among spicules from different parts of the sponge sample, suggesting that microhexactins are nearly absent in this species. 96 ZooKeys 1262: 81–96 (2025), DOI: 10.3897/zookeys.1262.164821 Lin Gong et al.: Two new hyalonematida species from the Indo-Pacific Supplementary material 3 trimmed alignments of 16S rDNA sequences of Hyalonema Authors: Lin Gong, Edwin Setiawan, Xinzheng Li, Swee Cheng Lim Explanation note: fas Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1262.164821.suppl3 Supplementary material 4 Phylogenetic tree obtained by maximum-likelihood (ML) analysis based on 16S rDNA Authors: Lin Gong, Edwin Setiawan, Xinzheng Li, Swee Cheng Lim Data type: tif Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1262.164821.suppl4