Hernández-Díaz YQ et al. (2023), Integrative species delimitation in the common ...

ECB-ART-51751

PeerJ 2023 Jul 17;11:e15655. doi: 10.7717/peerj.15655.

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Integrative species delimitation in the common ophiuroid Ophiothrix angulata (Echinodermata: Ophiuroidea): insights from COI, ITS2, arm coloration, and geometric morphometrics.

Hernández-Díaz YQ , Solis F , Beltrán-López RG , Benítez HA , Díaz-Jaimes P , Paulay G .

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Ophiothrix angulata (Say, 1825) is one of the most common and well-known ophiuroids in the Western Atlantic, with a wide geographic and bathymetric range. The taxonomy of this species has been controversial for a century because of its high morphological variability. Here we integrate information from DNA sequence data, color patterns, and geometric morphometrics to assess species delimitation and geographic differentiation in O. angulata. We found three deeply divergent mtDNA-COI clades (K2P 17.0-27.9%). ITS2 nuclear gene and geometric morphometrics of dorsal and ventral arm plates differentiate one of these lineages, as do integrative species delineation analyses, making this a confirmed candidate species.

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	Figure 1. Sampling locations (see Table S1 for details), with the proportion of specimens per clade are given as pie charts.Clade 1 (green), Clade 2 (blue), and Clade 3 (pink). OS: off Steinhatchee, North of St. Petersburg and Cedar Key. NWSP, Northwest of St. Petersburg; NSP and SSP, North and South of St. Petersburg; NFK, North of Florida Keys; FK, Florida Keys; PB, Palm Beach; JA, near Jacksonville; LU, near Louisiana; TV, Tuxpan Veracruz; SV, Sistema Arrecifal Veracruzano and Monte Pio; CAC, Cayo Arcas area; CAN, Cayo Arenas; ALA, Alacranes Reef; MR, Mayan Riviera; MH, Mahahual; BE, Belize; NI, Nicaragua; CR, Costa Rica; PA, Panama; StM, St. Martin; CU, Curaçao; GU, Guadeloupe; FG, French Guiana. ©OpenStreetMap 2022.
	Figure 2. DAP and VAP landmarks.Landmarks type II (red) and semi-landmarks (yellow) used for geometric morphometrics to investigate variation in arm plates in Ophiothrix angulata. Numbers indicate landmark position. (A) Dorsal arm plate (DAP). (B) Ventral arm plate (VAP). Pe, Proximal edge; De, Distal edge. Photos credit: Y. Quetzalli Hernández-Díaz.
	Figure 3. MtDNA bayesian consensus tree.Bayesian consensus tree of COI sequences produced using the GTR+I+G model in MrBayes v.3.2.7a on XSEDE for Ophiothrix angulata and outgroups. Clade 1 (green), Clade 2 (blue), Clade 3 (pink), and Outgroup (gray). Bayesian posterior probabilities (above), followed by ML bootstrap support (below; 1,000 replicates), are indicated at nodes.
	Figure 4. MtDNA vs nrDNA trees.A comparison of mtDNA (COI) (left) and nrDNA (ITS2) (right) phylogenies from the BI analysis are displayed as cladograms for Ophiothrix angulata with Bayesian posterior probabilities displayed above and ML bootstrap (1,000 replicates) support displayed below the nodes.
	Figure 5. TCS haplotype network clades 1 and 2.Clade 1A (n = 42), Clade 1B (n = 26), Clade 1C (n = 3), Clade 2A (n = 42), and Clade 2B (n = 2). TCS haplotype network of COI sequences for two Ophiothrix angulata clades (632 bp). The number of specimens is superimposed onto the more abundant haplotypes. Northern Gulf of Mexico (green). Florida Keys (orange). Eastern Florida (vermilion). Southern Gulf of Mexico (grey). Campeche Bank (yellow). Western Caribbean (blue). Southwestern Caribbean (sky blue). Eastern Caribbean (pink).
	Figure 6. TCS haplotype network Clade 3 (n = 31).TCS haplotype network of COI sequences for one Ophiothrix angulata clade (632 pb). The number of specimens is superimposed onto the more abundant haplotypes. Northern Gulf of Mexico (green). Florida Keys (orange). Campeche Bank (yellow). Western Caribbean (blue). Eastern Caribbean (pink).
	Figure 7. Dorsal arm plate geometric morphometrics.Scatterplots showing shape variation along principal component axes. Clade 1A (green). Clade 1B (orange). Clade 2 (blue). Clade 3 (pink). Thin-plate spline deformation grids accompany each PC axis to show the specimens’ shape at their positive and negative ends; the arm plate consensus shape is gray. The bar graph depicts the percentage of variance explained by PC axes.
	Figure 8. Ventral arm plate geometric morphometrics.Scatterplots showing shape variation along principal component axes. Clade 1A (green). Clade 1B (orange). Clade 2 (blue). Clade 3 (pink). Thin-plate spline deformation grids accompany each PC axis to show the specimens’ shape at their positive and negative ends; the arm plate consensus shape is gray. The bar graph depicts the percentage of variance explained by PC axes.
	Figure 9. Species delimitation for Ophiothrix angulata clades based on molecular, morphological, and integrative approaches.Clade 1A (UF7632), Clade 1B (UF11608), Clade 2A (UF10248), and Clade 3 (UF10250). COI phylogeny on the left, representative specimen based on DAP and VAP consensus shape of major clades in the middle, species delineations on the right. Delineated species are represented by separate colors. iBPP results are based on (A) COI and arm color information (Seq+COL) and (B) COI and geometric morphometric data (Seq+GM). ND, no data. Photo credit: Invertebrate Zoology Collection, Florida Museum of Natural History, University of Florida.

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