## Acknowledgements

We are grateful to the Tree of Life Programme at the Wellcome Sanger Institute that sequenced and assembled the *Asterias rubens* genome, in partnership with the Elphick lab, for the 25 Genomes Project (https://www.sanger.ac.uk/collaboration/25-genomes-for-25-years/) and the Darwin Tree of Life Project (https://www.darwintreeoflife.org/). The *A. rubens* genome sequence data are publicly available via the NCBI accession number PRJEB33974 [44]. M.R.E. dedicates this paper to the memory of Michael C. Thorndyke (1946-2022).

## Authors' contributions

M.R.E., N.E.C., D.C.S., and L.A.Y.G. conceived the study. N.E.C., D.C.S., L.A.Y.G., M.Z., S.E.S., J.H.S., C.G.Z., A.M.J., and M.R.E. performed the experimental work and/or analyzed the experimental data. N.E.C., L.A.Y.G., M.d.R., O.M., and M.R.E. performed or supervised the computational analysis of the sequence data. N.E.C., L.A.Y.G., and M.R.E. wrote the paper with input from all co-authors. The authors read and approved the final manuscript.

## Funding

The research reported in this paper was funded by grants awarded by the Leverhulme Trust to M.R.E (RPG-2013-351; RPG-2016-353) and by the Bio-technology and Biological Sciences Research Council (BBSRC) to M.R.E. (BB/M001644/1) and to J.H.S. and A.M.J. (BB/M001032/1). N.E.C. and L.A.Y.G were supported by PhD studentships awarded by the Mexican Council of Science and Technology (CONACyT studentship no. 471864 and 418612, respectively) and Queen Mary University of London.

## Availability of data and materials

All datasets generated or analyzed during this study are included in this published article and the supplementary additional files. The *A. rubens* genome sequence data analyzed in this paper are publicly available via the NCBI accession number PRJEB33974 [44].

## Declarations

### Ethics approval and consent to participate

Not applicable.

### Consent for publication

Not applicable.

### Competing interests

The authors declare that they have no competing interests.

### Author details

¹Queen Mary University of London, School of Biological & Behavioural Sciences, London E1 4NS, UK. ²Present address: Institute of Medical and Biomedical Education, St George's University of London, Cranmer Terrace, London SW17 0RE, UK. ³Present Address: Living Systems Institute, University of Exeter, Exeter EX4 4QD, UK. ⁴Present Address: Neurobiology and Genetics, Theodor-Boveri-Institute, Biocenter, University of Würzburg, 97074 Würzburg, Germany. ⁵School of Life Sciences, University of Warwick, Coventry CV4 7AL, UK. ⁶Present address: Waters Corporation, Stamford Avenue, Altrincham Road, Wilmslow SK9 4AX, UK. ⁷Present address: School of Science, Engineering & Design, Stephenson Street, Teesside University, Middlesbrough TS1 3BX, TS1 3BA, Tees Valley, UK. ⁸Institut Pasteur, Université Paris Cité, Bioinformatics and Biostatistics Hub, 75015 Paris, France.

Received: 27 April 2022 Accepted: 9 August 2022

Published online: 24 August 2022

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27. Koziol U. Precursors of neuropeptides and peptide hormones in the genomes of tardigrades. Gen Comp Endocrinol. 2018;267:12.

28. Nguyen TV, Rotllant GE, Cummins SF, Elizur A, Ventura T. Insights into sexual maturation and reproduction in the Norway lobster (Nephrops norvegicus) via in silico prediction and characterization of neuropeptides and G protein-coupled receptors. Front Endocrinol. 2018;9:16.

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31. Semmens DC, Mirabeau O, Moghul I, Pancholi MR, Wurm Y, Elphick MR. Transcriptomic identification of starfish neuropeptide precursors yields new insights into neuropeptide evolution. Open Biol. 2016;6(2): 150224.

32. Zandawala M, Moghul I, Yañez Guerra LA, Delroisse J, Abylkassimova N, Hugall AF, O'Hara TD, Elphick MR. Discovery of novel representatives of bilaterian neuropeptide families and reconstruction of neuropeptide precursor evolution in ophiuroid echinoderms. Open Biol. 2017;7:170129.

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34. Elphick MR, Price DA, Lee TD, Thorndyke MC. The SALMFamides: a new family of neuropeptides isolated from an echinoderm. Proceedings Biol Sci/ Royal Soc. 1991;243(1307):121-7.

35. Elphick MR. SALMFamide salmagundi: the biology of a neuropeptide family in echinoderms. Gen Comp Endocrinol. 2014;205:23-35.

36. Semmens DC, Dane RE, Pancholi MR, Slade SE, Scrivens JH, Elphick MR. Discovery of a novel neurophysin-associated neuropeptide that triggers cardiac stomach contraction and retraction in starfish. J Exp Biol. 2013;216(Pt 21):4047-53.

37. Yañez-Guerra LA, Delroisse J, Barreiro-Iglesias A, Slade SE, Scrivens JH, Elphick MR. Discovery and functional characterization of a lugin type neuropeptide signalling system in a deuterostome. Sci Rep. 2018;8(1):7220.

38. Tinoco AB, Barreiro-Iglesias A, Yañez Guerra LA, Delroisse J, Zhang Y, Gunner EF, Zampronio CG, Jones AM, Egertová M, Elphick MR. Ancient role of sulfakinin/cholecystokinin-type signalling in inhibitory regulation of feeding processes revealed in an echinoderm. Elife. 2021;10:e65667.

39. Tian S, Zandawala M, Beets I, Baytemur E, Slade SE, Scrivens JH, Elphick MR. Urbilaterian origin of paralogous GnRH and corazonin neuropeptide signalling pathways. Sci Rep. 2016;6:28788.

40. Zhang Y, Yañez Guerra LA, Egertová M, Zampronio CG, Jones AM, Elphick MR. Molecular and functional characterization of somatostatin-type signalling in a deuterostome invertebrate. Open Biol. 2020;10(9):200172.

41. Tinoco AB, Semmens DC, Patching EC, Gunner EF, Egertová M, Elphick MR. Characterization of NGFFYamide signaling in starfish reveals roles in regulation of feeding behavior and locomotory systems. Front Endocrinol (Lausanne). 2018;9:507.

42. Yañez-Guerra LA, Zhong X, Moghul I, Butts T, Zampronio CG, Jones AM, Mirabeau O, Elphick MR. Echinoderms provide missing link in the evolution of PrP/sNPF-type neuropeptide signalling. Elife. 2020;9:e57640.

43. Odekunle EA, Semmens DC, Martynyuk N, Tinoco AB, Garewal AK, Patel RR, Blowes LM, Zandawala M, Delroisse J, Slade SE, et al. Ancient role of vasopressin/oxytocin-type neuropeptides as regulators of feeding revealed in an echinoderm. BMC Biol. 2019;17(1):60.

44. Asterias rubens genome sequence [https://www.ncbi.nlm.nih.gov/bioproject/PRJEB33974]

45. Cai W, Kim CH, Go HJ, Egertová M, Zampronio CG, Jones AM, Park NG, Elphick MR. Biochemical, anatomical, and pharmacological characterization of calcitonin-type neuropeptides in starfish: discovery of an ancient role as muscle relaxants. Front Neurosci. 2018;12:382.

46. Lin M, Egertová M, Zampronio CG, Jones AM, Elphick MR. Pedal peptide/orcokinin-type neuropeptide signaling in a deuterostome: the anatomy and pharmacology of starfish myorelaxant peptide in Asterias rubens. J Comp Neurol. 2017;525(18):3890-917.

47. Lin M, Mita M, Egertová M, Zampronio CG, Jones AM, Elphick MR. Cellular localization of relaxin-like gonad-stimulating peptide expression in

Asterias rubens: new insights into neurohormonal control of spawning in starfish. J Comp Neurol. 2017;525(7):1599-617.

48. Elphick MR. Neuropeptide signaling in echinoderms: from "Physiologic Activity of Nerve Extracts" to neuropepididomics and beyond. In: Saleud-din S, Lange AB, Orchard I, editors. Advances in Invertebrate (Neuro) Endocrinology A Collection of Reviews in the Post-Genomic Era: Volume 1: Phyla Other Than Arthropoda. New York: Apple Academic Press; 2020. p. 125-71.

49. Zhang Y, Yañez-Guerra LA, Tinoco AB, Escudero Castelán N, Egertová M, Elphick MR. Somatostatin-type and allatostatin-C-type neuropeptides are paralogous and have opposing myoregulatory roles in an echinoderm. Proc Natl Acad Sci U S A. 2022;119(7):e2113589119.

50. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst Biol. 2010;59(3):307-21.

51. Guindon S, Gascuel O. A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst Biol. 2003;52(5):696-704.

52. Kim DK, Yun S, Son GH, Hwang JI, Park CR, Kim JI, Kim K, Vaudry H, Seong JY. Coevolution of the spexin/galanin/kisspeptin family: Spexin activates galanin receptor type II and III. Endocrinology. 2014;155(5):1864-73.

53. Smith MK, Wang T, Suwansa-Ard S, Motti CA, Elizur A, Zhao M, Rowe M, Hall MR, Elphick MR, Cummins SF. The neuropeptidome of the Crown-of-Thorns starfish, Acanthaster planci. J Proteomics. 2017;165:61-8.

54. Suwansa-Ard S, Chaiyamoon A, Talarovicova A, Tinkul R, Tinkul Y, Poomtong T, Elphick MR, Cummins SF, Sobhon P. Transcriptomic discovery and comparative analysis of neuropeptide precursors in sea cucumbers (Holothuroidea). Peptides. 2018;99:231-40.

55. Chen M, Talarovicova A, Zheng Y, Storey KB, Elphick MR. Neuropeptide precursors and neuropeptides in the sea cucumber Apostichopus japonicus: a genomic, transcriptomic and proteomic analysis. Sci Rep. 2019;9(1):1-22.

56. Elphick MR, Reeve JR Jr, Burke RD, Thorndyke MC. Isolation of the neuropeptide SALMFamide-1 from starfish using a new antiserum. Peptides. 1991;12(3):455-9.

57. Elphick MR, Semmens DC, Blowes LM, Levine J, Lowe CJ, Arnone MI, Clark MS. Reconstructing SALMFamide neuropeptide precursor evolution in the phylum Echinodermata: ophiuroid and crinoid sequence data provide new insights. Front Endocrinol (Lausanne). 2015;6:2.

58. Rowe ML, Elphick MR. Discovery of a second SALMFamide gene in the sea urchin Strongylocentrotus purpuratus reveals that L-type and F-type SALMFamide neuropeptides coexist in an echinoderm species. Mar Genomics. 2010;3(2):91-7.

59. Chieu HD, Suwansa-Ard S, Wang T, Elizur A, Cummins SF. Identification of neuropeptides in the sea cucumber Holothuria leucospilota. Gen Comp Endocrinol. 2019;283:113229.

60. Menschaert G, Vandekerckhove TT, Baggerman G, Landuyt B, Sweedler JV, Schoofs L, Luyten W, Van Criekinge W. A hybrid, de novo based, genome-wide database search approach applied to the sea urchin neuropeptide-dome. J Proteome Res. 2010;9(2):990-6.

61. Mayorova TD, Tian S, Cai W, Semmens DC, Odekunle EA, Zandawala M, Badi Y, Rowe ML, Egertová M, Elphick MR. Localization of neuropeptide gene expression in larvae of an echinoderm, the starfish Asterias rubens. Front Neurosci. 2016;10:553.

62. Newman SJ, Elphick MR, Thorndyke MC. Tissue distribution of the SALMFamide neuropeptides S1 and S2 in the starfish Asterias rubens using novel monoclonal and polyclonal antibodies. I. Nervous and locomotory systems. Proc Biol Sci. 1995;261(1360):139-45

63. Newman SJ, Elphick MR, Thorndyke MC. Tissue distribution of the SALMFamide neuropeptides S1 and S2 in the starfish Asterias rubens using novel monoclonal and polyclonal antibodies. 2. Digestive system. Proc Biol Sci. 1995;261(1360):187-92.

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26. Cardoso JCR, Félix RC, Bjärnmark N, Power DM. Allatostatin-type A, kisspeptin and galanin GPCRs and putative ligands as candidate regulatory factors of mantle function. Mar Genomics. 2016;27:11.

27. Koziol U. Precursors of neuropeptides and peptide hormones in the genomes of tardigrades. Gen Comp Endocrinol. 2018;267:12.

28. Nguyen TV, Rotllant GE, Cummins SF, Elizur A, Ventura T. Insights into sexual maturation and reproduction in the Norway lobster (Nephrops norvegicus) via in silico prediction and characterization of neuropeptides and G protein-coupled receptors. Front Endocrinol. 2018;9:16.

29. Veenstra JA. Similarities between decapod and insect neuropeptidomes. PeerJ. 2016;4(e2043):38.

30. Veenstra JA. Coleoptera genome and transcriptome sequences reveal numerous differences in neuropeptide signaling between species. PeerJ. 2019;7(e7144):39.

31. Semmens DC, Mirabeau O, Moghul I, Pancholi MR, Wurm Y, Elphick MR. Transcriptomic identification of starfish neuropeptide precursors yields new insights into neuropeptide evolution. Open Biol. 2016;6(2): 150224.

32. Zandawala M, Moghul I, Yañez Guerra LA, Delroisse J, Abylkassimova N, Hugall AF, O'Hara TD, Elphick MR. Discovery of novel representatives of bilaterian neuropeptide families and reconstruction of neuropeptide precursor evolution in ophiuroid echinoderms. Open Biol. 2017;7:170129.

33. Wang T, Cao Z, Shen Z, Yang J, Chen X, Yang Z, Xu K, Xiang X, Yu Q, Song Y, et al. Existence and functions of a kisspeptin neuropeptide signaling system in a non-chordate deuterostome species. Elife. 2020;9:e53370.

34. Elphick MR, Price DA, Lee TD, Thorndyke MC. The SALMFamides: a new family of neuropeptides isolated from an echinoderm. Proceedings Biol Sci/ Royal Soc. 1991;243(1307):121-7.

35. Elphick MR. SALMFamide salmagundi: the biology of a neuropeptide family in echinoderms. Gen Comp Endocrinol. 2014;205:23-35.

36. Semmens DC, Dane RE, Pancholi MR, Slade SE, Scrivens JH, Elphick MR. Discovery of a novel neurophysin-associated neuropeptide that triggers cardiac stomach contraction and retraction in starfish. J Exp Biol. 2013;216(Pt 21):4047-53.

37. Yañez-Guerra LA, Delroisse J, Barreiro-Iglesias A, Slade SE, Scrivens JH, Elphick MR. Discovery and functional characterization of a lugin type neuropeptide signalling system in a deuterostome. Sci Rep. 2018;8(1):7220.

38. Tinoco AB, Barreiro-Iglesias A, Yañez Guerra LA, Delroisse J, Zhang Y, Gunner EF, Zampronio CG, Jones AM, Egertová M, Elphick MR. Ancient role of sulfakinin/cholecystokinin-type signalling in inhibitory regulation of feeding processes revealed in an echinoderm. Elife. 2021;10:e65667.

39. Tian S, Zandawala M, Beets I, Baytemur E, Slade SE, Scrivens JH, Elphick MR. Urbilaterian origin of paralogous GnRH and corazonin neuropeptide signalling pathways. Sci Rep. 2016;6:28788.

40. Zhang Y, Yañez Guerra LA, Egertová M, Zampronio CG, Jones AM, Elphick MR. Molecular and functional characterization of somatostatin-type signalling in a deuterostome invertebrate. Open Biol. 2020;10(9):200172.

41. Tinoco AB, Semmens DC, Patching EC, Gunner EF, Egertová M, Elphick MR. Characterization of NGFFYamide signaling in starfish reveals roles in regulation of feeding behavior and locomotory systems. Front Endocrinol (Lausanne). 2018;9:507.

42. Yañez-Guerra LA, Zhong X, Moghul I, Butts T, Zampronio CG, Jones AM, Mirabeau O, Elphick MR. Echinoderms provide missing link in the evolution of PrP/sNPF-type neuropeptide signalling. Elife. 2020;9:e57640.

43. Odekunle EA, Semmens DC, Martynyuk N, Tinoco AB, Garewal AK, Patel RR, Blowes LM, Zandawala M, Delroisse J, Slade SE, et al. Ancient role of vasopressin/oxytocin-type neuropeptides as regulators of feeding revealed in an echinoderm. BMC Biol. 2019;17(1):60.

44. Asterias rubens genome sequence [https://www.ncbi.nlm.nih.gov/bioproject/PRJEB33974]

45. Cai W, Kim CH, Go HJ, Egertová M, Zampronio CG, Jones AM, Park NG, Elphick MR. Biochemical, anatomical, and pharmacological characterization of calcitonin-type neuropeptides in starfish: discovery of an ancient role as muscle relaxants. Front Neurosci. 2018;12:382.

46. Lin M, Egertová M, Zampronio CG, Jones AM, Elphick MR. Pedal peptide/orcokinin-type neuropeptide signaling in a deuterostome: the anatomy and pharmacology of starfish myorelaxant peptide in Asterias rubens. J Comp Neurol. 2017;525(18):3890-917.

47. Lin M, Mita M, Egertová M, Zampronio CG, Jones AM, Elphick MR. Cellular localization of relaxin-like gonad-stimulating peptide expression in

Asterias rubens: new insights into neurohormonal control of spawning in starfish. J Comp Neurol. 2017;525(7):1599-617.

48. Elphick MR. Neuropeptide signaling in echinoderms: from "Physiologic Activity of Nerve Extracts" to neuropepididomics and beyond. In: Saleud-din S, Lange AB, Orchard I, editors. Advances in Invertebrate (Neuro) Endocrinology A Collection of Reviews in the Post-Genomic Era: Volume 1: Phyla Other Than Arthropoda. New York: Apple Academic Press; 2020. p. 125-71.

49. Zhang Y, Yañez-Guerra LA, Tinoco AB, Escudero Castelán N, Egertová M, Elphick MR. Somatostatin-type and allatostatin-C-type neuropeptides are paralogous and have opposing myoregulatory roles in an echinoderm. Proc Natl Acad Sci U S A. 2022;119(7):e2113589119.

50. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst Biol. 2010;59(3):307-21.

51. Guindon S, Gascuel O. A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst Biol. 2003;52(5):696-704.

52. Kim DK, Yun S, Son GH, Hwang JI, Park CR, Kim JI, Kim K, Vaudry H, Seong JY. Coevolution of the spexin/galanin/kisspeptin family: Spexin activates galanin receptor type II and III. Endocrinology. 2014;155(5):1864-73.

53. Smith MK, Wang T, Suwansa-Ard S, Motti CA, Elizur A, Zhao M, Rowe M, Hall MR, Elphick MR, Cummins SF. The neuropeptidome of the Crown-of-Thorns starfish, Acanthaster planci. J Proteomics. 2017;165:61-8.

54. Suwansa-Ard S, Chaiyamoon A, Talarovicova A, Tinkul R, Tinkul Y, Poomtong T, Elphick MR, Cummins SF, Sobhon P. Transcriptomic discovery and comparative analysis of neuropeptide precursors in sea cucumbers (Holothuroidea). Peptides. 2018;99:231-40.

55. Chen M, Talarovicova A, Zheng Y, Storey KB, Elphick MR. Neuropeptide precursors and neuropeptides in the sea cucumber Apostichopus japonicus: a genomic, transcriptomic and proteomic analysis. Sci Rep. 2019;9(1):1-22.

56. Elphick MR, Reeve JR Jr, Burke RD, Thorndyke MC. Isolation of the neuropeptide SALMFamide-1 from starfish using a new antiserum. Peptides. 1991;12(3):455-9.

57. Elphick MR, Semmens DC, Blowes LM, Levine J, Lowe CJ, Arnone MI, Clark MS. Reconstructing SALMFamide neuropeptide precursor evolution in the phylum Echinodermata: ophiuroid and crinoid sequence data provide new insights. Front Endocrinol (Lausanne). 2015;6:2.

58. Rowe ML, Elphick MR. Discovery of a second SALMFamide gene in the sea urchin Strongylocentrotus purpuratus reveals that L-type and F-type SALMFamide neuropeptides coexist in an echinoderm species. Mar Genomics. 2010;3(2):91-7.

59. Chieu HD, Suwansa-Ard S, Wang T, Elizur A, Cummins SF. Identification of neuropeptides in the sea cucumber Holothuria leucospilota. Gen Comp Endocrinol. 2019;283:113229.

60. Menschaert G, Vandekerckhove TT, Baggerman G, Landuyt B, Sweedler JV, Schoofs L, Luyten W, Van Criekinge W. A hybrid, de novo based, genome-wide database search approach applied to the sea urchin neuropeptide-dome. J Proteome Res. 2010;9(2):990-6.

61. Mayorova TD, Tian S, Cai W, Semmens DC, Odekunle EA, Zandawala M, Badi Y, Rowe ML, Egertová M, Elphick MR. Localization of neuropeptide gene expression in larvae of an echinoderm, the starfish Asterias rubens. Front Neurosci. 2016;10:553.

62. Newman SJ, Elphick MR, Thorndyke MC. Tissue distribution of the SALMFamide neuropeptides S1 and S2 in the starfish Asterias rubens using novel monoclonal and polyclonal antibodies. I. Nervous and locomotory systems. Proc Biol Sci. 1995;261(1360):139-45

63. Newman SJ, Elphick MR, Thorndyke MC. Tissue distribution of the SALMFamide neuropeptides S1 and S2 in the starfish Asterias rubens using novel monoclonal and polyclonal antibodies. 2. Digestive system. Proc Biol Sci. 1995;261(1360):187-92.

64. Moss C, Burke RD, Thorndyke MC. Immunocytochemical localization of the neuropeptide S1 and serotonin in larvae of the starfish Pisaster ochraceus and Asterias rubens. J Mar Biol Assoc UK. 1994;74(1):61-71.

65. Elphick MR, Newman SJ, Thorndyke MC. Distribution and action of SLAMFamide neuropeptides in the starfish Asterias rubens. J Exp Biol. 1995;198(12):2519-25.

66. Melange R, Potton DJ, Thorndyke MC, Elphick MR. SALMFamide neuropeptides cause relaxation and eversion of the cardiac stomach in starfish. Proc Biol Sci. 1999;266:1785-9.
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68. Mita M, Oka H, Thorndyke MC, Shibata Y, Yoshikuni M, Nagahama Y. Inhibitory effect of a SALMFamide neuropeptide on secretion of gonad-stimulating substance from radial nerves in the starfish Asterina pectinifera. Zoolog Sci. 2004;21(3):299-303.

69. Li W, Godzik A. Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics Applications Note. 2006;22(13):1658-9.

70. Chen C, Chen H, Zhang Y, Thomas HR, Frank MH, He Y, Xia R. TBtools: an integrative toolkit developed for interactive analyses of big biological data. Mol Plant. 2020;13(8):1194-202.

71. Ranwez V, Gascuel O. Improvement of distance-based phylogenetic methods by a local maximum likelihood approach using triplets. Mol Biol Evol. 2002;19(11):12.

72. Capella-Gutierrez S, Silla-Martinez JM, Gabaldon T. trimAI: a tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics. 2009;25(15):1972-3.

73. Minh BQ, Schmidt HA, Chernomor O, Scherempf D, Woodhams MD, Von Haeseler A, Lanfear R. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Mol Biol Evol. 2020;37(5):1530-4.

74. Nguyen L-T, Schmidt HA, Von Haeseler A, Minh BQ. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol Evol. 2015;32(1):268-74.

75. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology. 2010;59(3):307-21.

76. Hoang DT, Chernomor O, Von Haeseler A, Minh BQ, Vinh LS. UFBoot2: improving the ultrafast bootstrap approximation. Mol Biol Evol. 2018;35(2):518-22.

77. Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchards MA. Hue;esenbeck JP: MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology. 2012;61(3):539-42.

78. Ayres DL, Darling A, Zwickl DJ, Beerli P, Holder M, Lewis PO, Huelsenbeck JP, Ronquist F, Swofford DL, Cummings MP, et al. BEAGLE: an application programming interface and high-performance computing library for statistical phylogenetics. Syst Biol. 2012;61(1):170-3.

79. Frickey T, Lupas A. CLANS: a Java application for visualizing protein families based on pairwise similarity. Bioinformatics. 2004;20(18):3702-4.

80. Kiryutin B, Souvorov A, Tatusova T: ProSplin-protein to genomic alignment tool. In: In Proc 11th Annual International Conference in Research in Computational Molecular Biology: 2007. 2.

81. Liu W, Xie Y, Ma J, Luo X, Nie P, Zuo Z, Lahrmann U, Zhao Q, Zheng Y, Zhao Y, et al. IBS: an illustrator for the presentation and visualization of biological sequences. Bioinformatics. 2015;31(20):3359-61.

82. Krzywinski M, Schein J, Birol I, Connors J, Gascoyne R, Horsman D, Jones SJ, Marra MA. Circos: an information aesthetic for comparative genomics. Genome Res. 2009;19(9):8.

83. Baubet V, Le Mouellic H, Campbell AK, Lucas-Meunier E, Fossier P, Brûlet P. Chimeric green fluorescent protein-aequorin as bioluminescent Ca2 reporters at the single-cell level. Proc Natl Acad Sci USA. 2000;97(12):6.

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<figure><img src="image_7.png" /></figure>
67. Díaz-Miranda L, García-Arrarás JE. Pharmacological action of the hepta-peptide GFSKLYFamide in the muscle of the sea cucumber Holothuria glaberrima (Echinodermata). Comp Biochem Physiol C Pharmacol Toxicol Endocrinol. 1995;110(2):171-6.

68. Mita M, Oka H, Thorndyke MC, Shibata Y, Yoshikuni M, Nagahama Y. Inhibitory effect of a SALMFamide neuropeptide on secretion of gonad-stimulating substance from radial nerves in the starfish Asterina pectinifera. Zoolog Sci. 2004;21(3):299-303.

69. Li W, Godzik A. Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics Applications Note. 2006;22(13):1658-9.

70. Chen C, Chen H, Zhang Y, Thomas HR, Frank MH, He Y, Xia R. TBtools: an integrative toolkit developed for interactive analyses of big biological data. Mol Plant. 2020;13(8):1194-202.

71. Ranwez V, Gascuel O. Improvement of distance-based phylogenetic methods by a local maximum likelihood approach using triplets. Mol Biol Evol. 2002;19(11):12.

72. Capella-Gutierrez S, Silla-Martinez JM, Gabaldon T. trimAI: a tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics. 2009;25(15):1972-3.

73. Minh BQ, Schmidt HA, Chernomor O, Scherempf D, Woodhams MD, Von Haeseler A, Lanfear R. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Mol Biol Evol. 2020;37(5):1530-4.

74. Nguyen L-T, Schmidt HA, Von Haeseler A, Minh BQ. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol Evol. 2015;32(1):268-74.

75. Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology. 2010;59(3):307-21.

76. Hoang DT, Chernomor O, Von Haeseler A, Minh BQ, Vinh LS. UFBoot2: improving the ultrafast bootstrap approximation. Mol Biol Evol. 2018;35(2):518-22.

77. Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchards MA. Hue;esenbeck JP: MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology. 2012;61(3):539-42.

78. Ayres DL, Darling A, Zwickl DJ, Beerli P, Holder M, Lewis PO, Huelsenbeck JP, Ronquist F, Swofford DL, Cummings MP, et al. BEAGLE: an application programming interface and high-performance computing library for statistical phylogenetics. Syst Biol. 2012;61(1):170-3.

79. Frickey T, Lupas A. CLANS: a Java application for visualizing protein families based on pairwise similarity. Bioinformatics. 2004;20(18):3702-4.

80. Kiryutin B, Souvorov A, Tatusova T: ProSplin-protein to genomic alignment tool. In: In Proc 11th Annual International Conference in Research in Computational Molecular Biology: 2007. 2.

81. Liu W, Xie Y, Ma J, Luo X, Nie P, Zuo Z, Lahrmann U, Zhao Q, Zheng Y, Zhao Y, et al. IBS: an illustrator for the presentation and visualization of biological sequences. Bioinformatics. 2015;31(20):3359-61.

82. Krzywinski M, Schein J, Birol I, Connors J, Gascoyne R, Horsman D, Jones SJ, Marra MA. Circos: an information aesthetic for comparative genomics. Genome Res. 2009;19(9):8.

83. Baubet V, Le Mouellic H, Campbell AK, Lucas-Meunier E, Fossier P, Brûlet P. Chimeric green fluorescent protein-aequorin as bioluminescent Ca2 reporters at the single-cell level. Proc Natl Acad Sci USA. 2000;97(12):6.

## Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Ready to submit your research? Choose BMC and benefit from:

* fast, convenient online submission

* thorough peer review by experienced researchers in your field

* rapid publication on acceptance

* support for research data, including large and complex data types

* gold Open Access which fosters wider collaboration and increased citations

* maximum visibility for your research: over 100M website views per year

At BMC, research is always in progress.

Learn more biomedcentral.com/submissions

<figure><img src="image_7.png" /></figure>