Project 4138: B. Bomfleur, G. W. Grimm, S. McLoughlin. 2017. The fossil Osmundales (Royal Ferns)—a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes. PeerJ. 5:e3433.
Abstract
The Osmundales (Royal Fern order) originated in the late Paleozoic and is the most ancient surviving lineage of leptosporangiate ferns. In contrast to its low diversity today (less than 20 species in six genera), it has the richest fossil record of any extant group of ferns. The structurally preserved trunks and rhizomes alone are referable to more than 100 fossil species that are classified in up to 20 genera, four subfamilies, and two families. This diverse fossil record constitutes an exceptional source of information on the evolutionary history of the group from the Permian to the present. However, inconsistent terminology, varying formats of description, and the general lack of a uniform taxonomic concept renders this wealth of information poorly accessible. To this end, we provide a comprehensive review of the diversity of structural features of osmundalean axes under a standardized, descriptive terminology. A novel morphological character matrix with 45 anatomical characters scored for 15 extant species and for 114 fossil operational units (species or specimens) is analysed using networks in order to establish systematic relationships among fossil and extant Osmundales rooted in axis anatomy. The results lead us to propose an evolutionary classification for fossil Osmundales and a revised, standardized taxonomy for all taxa down to the rank of (sub)genus. We introduce several nomenclatural novelties: (1) a new subfamily Itopsidemoideae (Guaireaceae) is established to contain Itopsidema, Donwelliacaulis, and Tiania; (2) the thamnopteroid genera Zalesskya, Iegosigopteris, and Petcheropteris are all considered synonymous with Thamnopteris; (3) 12 species of Millerocaulis and Ashicaulis are assigned to modern genera (tribe Osmundeae); (4) the hitherto enigmatic Aurealcaulis is identified as an extinct subgenus of Plenasium; and (5) the poorly known Osmundites tuhajkulensis is assigned to Millerocaulis. In addition, we consider Millerocaulis stipabonettiorum a possible member of Palaeosmunda and Millerocaulis estipularis as probably constituting the earliest representative of the (Todea-)Leptopteris lineage (subtribe Todeinae) of modern Osmundoideae.Read the article »
Article DOI: 10.7717/peerj.3433
Project DOI: 10.7934/P4138, http://dx.doi.org/10.7934/P4138
| This project contains | Matrices |
|---|---|
Download Project SDD File  | Total scored cells: 4823 Total media associated with cells: 0 Total labels associated with cell media: 0 |
| Characters | |
| Total characters: 45 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 112 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:45/0 |
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MorphoBank Project 4138
MorphoBank Project 4138
- Creation Date:
01 December 2021 - Publication Date:
29 January 2022 - Project views: 6645
- Matrix downloads: 1
Authors' Institutions
- Swedish Museum of Natural History (Naturhistoriska riksmuseet) Stockholm
- Universitaet Muenster (Westfälische Wilhelms-Universität Münster)
- University of Vienna
Members
| member name | taxa |
specimens |
media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
| MorphoBank Curator Project Administrator | 131 | 1 | 1 | 45 | 0 | 0 | 4823 (4823, 0, 0) | 0 | 0 | 0 | ||||
| Benjamin Bomfleur Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
| MorphoBank Curator Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Taxonomic Overview for Matrix 'M27717' (131 Taxa)
| taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
| [1] † Anomorrhoea fischeri Last Modified in 12/03/21 | 31 | 14 | 14 | 0 | 0 | 0 | 0 | 3 |
| [2] † Bathypteris rhomboidea Last Modified in 12/03/21 | 17 | 29 | 28 | 0 | 0 | 0 | 0 | 3 |
| [3] † Chasmatopteris principalis Last Modified in 12/03/21 | 12 | 33 | 32 | 0 | 0 | 0 | 0 | 3 |
| [4] † Grammatopteris baldaufii Last Modified in 12/03/21 | 15 | 33 | 30 | 0 | 0 | 0 | 0 | 3 |
| [5] † Grammatopteris rigollotii Last Modified in 12/03/21 | 16 | 29 | 29 | 0 | 0 | 0 | 0 | 3 |
| [6] † Grammatopteris freitasii Last Modified in 12/03/21 | 15 | 31 | 30 | 0 | 0 | 0 | 0 | 3 |
| [7] † Guairea brasiliensis Last Modified in 12/03/21 | 21 | 27 | 24 | 0 | 0 | 0 | 0 | 3 |
| [8] † Guairea carnierii Last Modified in 12/03/21 | 21 | 27 | 24 | 0 | 0 | 0 | 0 | 3 |
| [9] † Guairea millerii Last Modified in 12/03/21 | 20 | 26 | 25 | 0 | 0 | 0 | 0 | 3 |
| [10] † Palaeosmunda playfordii Last Modified in 12/03/21 | 8 | 38 | 36 | 0 | 0 | 0 | 0 | 3 |
| [11] † Palaeosmunda williamsii Last Modified in 12/03/21 | 6 | 42 | 38 | 0 | 0 | 0 | 0 | 3 |
| [12] † Rastropteris pinquanensis Last Modified in 12/03/21 | 16 | 29 | 29 | 0 | 0 | 0 | 0 | 3 |
| [13] † Shuichengella primitiva Last Modified in 12/03/21 | 19 | 26 | 26 | 0 | 0 | 0 | 0 | 3 |
| [14] † Thamnopteris diploxylon Last Modified in 12/03/21 | 25 | 22 | 20 | 0 | 0 | 0 | 0 | 3 |
| [15] † Thamnopteris gracilis Last Modified in 12/03/21 | 24 | 23 | 21 | 0 | 0 | 0 | 0 | 3 |
| [16] † Thamnopteris gwynnevaughanii Last Modified in 12/03/21 | 12 | 35 | 33 | 0 | 0 | 0 | 0 | 3 |
| [17] † Thamnopteris javorskii Last Modified in 12/03/21 | 10 | 38 | 32 | 0 | 0 | 0 | 0 | 3 |
| [18] † Thamnopteris kazanensis Last Modified in 12/03/21 | 25 | 22 | 20 | 0 | 0 | 0 | 0 | 3 |
| [19] † Thamnopteris kidstonii Last Modified in 12/03/21 | 7 | 42 | 38 | 0 | 0 | 0 | 0 | 3 |
| [20] † Thamnopteris schlechtendalii Last Modified in 12/03/21 | 9 | 38 | 36 | 0 | 0 | 0 | 0 | 3 |
| [21] † Thamnopteris splendida Last Modified in 12/03/21 | 13 | 32 | 32 | 0 | 0 | 0 | 0 | 3 |
| [22] † Thamnopteris uralica Last Modified in 12/03/21 | 24 | 23 | 21 | 0 | 0 | 0 | 0 | 3 |
| [23] † Tiania yunnanensis Last Modified in 12/03/21 | 20 | 26 | 25 | 0 | 0 | 0 | 0 | 3 |
| [24] † Zhongmingella plenasioides Last Modified in 12/03/21 | 24 | 23 | 21 | 0 | 0 | 0 | 0 | 3 |
| [25] † Osmunda beardmorensis Last Modified in 12/03/21 | 7 | 41 | 38 | 0 | 0 | 0 | 0 | 3 |
| [26] † Millerocaulis herbstii Last Modified in 12/03/21 | 17 | 29 | 28 | 0 | 0 | 0 | 0 | 3 |
| [27] † Millerocaulis lutziae Last Modified in 12/03/21 | 18 | 27 | 27 | 0 | 0 | 0 | 0 | 3 |
| [28] † Millerocaulis woolfei Last Modified in 12/03/21 | 7 | 39 | 38 | 0 | 0 | 0 | 0 | 3 |
| [29] † Donwelliacaulis chlouberii Last Modified in 12/03/21 | 19 | 26 | 26 | 0 | 0 | 0 | 0 | 3 |
| [30] † Itopsidema vancleavei Last Modified in 12/03/21 | 18 | 27 | 27 | 0 | 0 | 0 | 0 | 3 |
| [31] † Osmunda indentata Last Modified in 12/03/21 | 3 | 42 | 42 | 0 | 0 | 0 | 0 | 3 |
| [32] † Millerocaulis stipabonettiorum Last Modified in 12/03/21 | 18 | 27 | 27 | 0 | 0 | 0 | 0 | 3 |
| [33] † Millerocaulis tuhajkulensis Last Modified in 12/03/21 | 17 | 28 | 27 | 0 | 0 | 0 | 0 | 3 |
| [34] † Millerocaulis aucklandicus Last Modified in 12/03/21 | 7 | 40 | 37 | 0 | 0 | 0 | 0 | 3 |
| [35] † Millerocaulis beipiaoensis Last Modified in 12/03/21 | 7 | 38 | 38 | 0 | 0 | 0 | 0 | 3 |
| [36] † Millerocaulis broganii Last Modified in 12/03/21 | 5 | 40 | 40 | 0 | 0 | 0 | 0 | 3 |
| [37] † Millerocaulis gibbianus Last Modified in 12/03/21 | 6 | 40 | 39 | 0 | 0 | 0 | 0 | 3 |
| [38] † Millerocaulis hebeiensis Last Modified in 12/03/21 | 8 | 37 | 37 | 0 | 0 | 0 | 0 | 3 |
| [39] † Millerocaulis macromedullosus Last Modified in 12/03/21 | 9 | 36 | 36 | 0 | 0 | 0 | 0 | 3 |
| [40] † Millerocaulis patagonicus Last Modified in 12/03/21 | 7 | 39 | 38 | 0 | 0 | 0 | 0 | 3 |
| [41] † Millerocaulis richmondii Last Modified in 12/03/21 | 5 | 40 | 40 | 0 | 0 | 0 | 0 | 3 |
| [42] † Millerocaulis santaecrucis Last Modified in 12/03/21 | 14 | 31 | 31 | 0 | 0 | 0 | 0 | 3 |
| [43] † Millerocaulis spinksii Last Modified in 12/03/21 | 8 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [44] † Millerocaulis swanensis Last Modified in 12/03/21 | 5 | 41 | 40 | 0 | 0 | 0 | 0 | 3 |
| [45] † Millerocaulis wadei Last Modified in 12/03/21 | 6 | 42 | 39 | 0 | 0 | 0 | 0 | 3 |
| [46] † Millerocaulis websteri Last Modified in 12/03/21 | 7 | 39 | 38 | 0 | 0 | 0 | 0 | 3 |
| [47] † Millerocaulis wrightii Last Modified in 12/03/21 | 7 | 39 | 38 | 0 | 0 | 0 | 0 | 3 |
| [48] † Grammatocaulis donponii Last Modified in 12/03/21 | 12 | 33 | 32 | 0 | 0 | 0 | 0 | 3 |
| [49] † Lunea jonesii Last Modified in 12/03/21 | 10 | 35 | 35 | 0 | 0 | 0 | 0 | 3 |
| [50] † Millerocaulis chubutensis Last Modified in 12/03/21 | 10 | 35 | 35 | 0 | 0 | 0 | 0 | 3 |
| [51] † Millerocaulis donponii Last Modified in 12/03/21 | 9 | 36 | 36 | 0 | 0 | 0 | 0 | 3 |
| [52] † Millerocaulis dunlopii Last Modified in 12/03/21 | 10 | 36 | 35 | 0 | 0 | 0 | 0 | 3 |
| [53] † Millerocaulis juandahensis Last Modified in 12/03/21 | 9 | 36 | 35 | 0 | 0 | 0 | 0 | 3 |
| [54] † Millerocaulis limewoodensis Last Modified in 12/03/21 | 9 | 36 | 36 | 0 | 0 | 0 | 0 | 3 |
| [55] † Osmunda preosmunda Last Modified in 12/03/21 | 5 | 40 | 40 | 0 | 0 | 0 | 0 | 3 |
| [56] † Osmunda sinica Last Modified in 12/03/21 | 3 | 42 | 42 | 0 | 0 | 0 | 0 | 3 |
| [57] † Osmunda chengii Last Modified in 12/03/21 | 1 | 44 | 44 | 0 | 0 | 0 | 0 | 3 |
| [58] † Osmunda liaoningensis Last Modified in 12/03/21 | 1 | 47 | 44 | 0 | 0 | 0 | 0 | 3 |
| [59] † Osmunda pulchella Last Modified in 12/03/21 | 2 | 45 | 43 | 0 | 0 | 0 | 0 | 3 |
| [60] † Osmunda plumites Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [61] † Osmunda wangii Last Modified in 12/03/21 | 2 | 44 | 43 | 0 | 0 | 0 | 0 | 3 |
| [62] † Osmundacaulis andrewii Last Modified in 12/03/21 | 7 | 40 | 38 | 0 | 0 | 0 | 0 | 3 |
| [63] † Osmundacaulis griggsii Last Modified in 12/03/21 | 5 | 43 | 40 | 0 | 0 | 0 | 0 | 3 |
| [64] † Osmundacaulis hoskingii var hoskingii Last Modified in 12/03/21 | 6 | 39 | 39 | 0 | 0 | 0 | 0 | 3 |
| [65] † Osmundacaulis janae Last Modified in 12/03/21 | 30 | 16 | 15 | 0 | 0 | 0 | 0 | 3 |
| [66] † Osmundacaulis jonesii Last Modified in 12/03/21 | 5 | 41 | 40 | 0 | 0 | 0 | 0 | 3 |
| [67] † Osmundacaulis lemonii Last Modified in 12/03/21 | 8 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [68] † Osmundacaulis nerii Last Modified in 12/03/21 | 6 | 41 | 39 | 0 | 0 | 0 | 0 | 3 |
| [69] † Osmundacaulis pruchnickii Last Modified in 12/03/21 | 5 | 42 | 40 | 0 | 0 | 0 | 0 | 3 |
| [70] † Osmundacaulis richmondii Last Modified in 12/03/21 | 6 | 40 | 39 | 0 | 0 | 0 | 0 | 3 |
| [71] † Osmundacaulis tasmanensis Last Modified in 12/03/21 | 7 | 38 | 38 | 0 | 0 | 0 | 0 | 3 |
| [72] † Osmundacaulis tehuelchensis Last Modified in 12/03/21 | 6 | 41 | 39 | 0 | 0 | 0 | 0 | 3 |
| [73] † Osmunda johnstonii Last Modified in 12/03/21 | 1 | 44 | 44 | 0 | 0 | 0 | 0 | 3 |
| [74] † Osmundacaulis bamfordiae Last Modified in 12/03/21 | 11 | 34 | 34 | 0 | 0 | 0 | 0 | 3 |
| [75] † Osmundacaulis tidwellii Last Modified in 12/03/21 | 9 | 37 | 36 | 0 | 0 | 0 | 0 | 3 |
| [76] † Osmundacaulis zululandensis Last Modified in 12/03/21 | 8 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [77] † Gen indet estipularis Last Modified in 12/03/21 | 13 | 32 | 30 | 0 | 0 | 0 | 0 | 3 |
| [78] † Millerocaulis amarjolensis Last Modified in 12/03/21 | 7 | 38 | 38 | 0 | 0 | 0 | 0 | 3 |
| [79] † Millerocaulis australis Last Modified in 12/03/21 | 8 | 37 | 37 | 0 | 0 | 0 | 0 | 3 |
| [80] † Millerocaulis indica Last Modified in 12/03/21 | 11 | 34 | 33 | 0 | 0 | 0 | 0 | 3 |
| [81] † Millerocaulis guptai Last Modified in 12/03/21 | 10 | 35 | 35 | 0 | 0 | 0 | 0 | 3 |
| [82] † Millerocaulis kolbei Last Modified in 12/03/21 | 7 | 40 | 37 | 0 | 0 | 0 | 0 | 3 |
| [83] † Millerocaulis livingstonensis Last Modified in 12/03/21 | 7 | 38 | 38 | 0 | 0 | 0 | 0 | 3 |
| [84] † Millerocaulis rajmahalensis Last Modified in 12/03/21 | 5 | 40 | 40 | 0 | 0 | 0 | 0 | 3 |
| [85] † Millerocaulis sahnii Last Modified in 12/03/21 | 8 | 37 | 37 | 0 | 0 | 0 | 0 | 3 |
| [86] † Osmunda tekelili Last Modified in 12/03/21 | 6 | 40 | 39 | 0 | 0 | 0 | 0 | 3 |
| [87] † Osmunda burgii Last Modified in 12/03/21 | 5 | 40 | 38 | 0 | 0 | 0 | 0 | 3 |
| [88] † Osmunda dakotensis Last Modified in 12/03/21 | 7 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [89] † Osmunda nebraskensis Last Modified in 12/03/21 | 7 | 39 | 38 | 0 | 0 | 0 | 0 | 3 |
| [90] † Osmunda embreei Last Modified in 12/03/21 | 3 | 44 | 42 | 0 | 0 | 0 | 0 | 3 |
| [91] † Osmunda cinnamomea Cretaceous Canada Last Modified in 12/03/21 | 2 | 43 | 42 | 0 | 0 | 0 | 0 | 3 |
| [92] † Osmunda kidstonii Last Modified in 12/03/21 | 31 | 14 | 14 | 0 | 0 | 0 | 0 | 3 |
| [93] † Osmundacaulis atherstonei Last Modified in 12/03/21 | 10 | 37 | 35 | 0 | 0 | 0 | 0 | 3 |
| [94] † Osmundacaulis natalensis Last Modified in 12/03/21 | 9 | 38 | 36 | 0 | 0 | 0 | 0 | 3 |
| [95] † Osmundacaulis skidegatensis Last Modified in 12/03/21 | 6 | 41 | 39 | 0 | 0 | 0 | 0 | 3 |
| [96] † Osmundacaulis whittlesii Last Modified in 12/03/21 | 8 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [97] † Todea tidwellii Last Modified in 12/03/21 | 11 | 34 | 33 | 0 | 0 | 0 | 0 | 3 |
| [98] † Osmunda bransonii Last Modified in 12/03/21 | 8 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [99] † Osmunda moorei Last Modified in 12/03/21 | 6 | 39 | 37 | 0 | 0 | 0 | 0 | 3 |
| [100] † Osmunda crossii Last Modified in 12/03/21 | 8 | 38 | 33 | 0 | 0 | 0 | 0 | 3 |
| [101] † Osmunda arnoldii Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [102] † ALT1 osmunda dowkeri MILLER Last Modified in 12/03/21 | 3 | 43 | 42 | 0 | 0 | 0 | 0 | 3 |
| [103] † ALT2 osmunda dowkeri CHANDLER Last Modified in 12/03/21 | 2 | 44 | 42 | 0 | 0 | 0 | 0 | 3 |
| [104] † ALT2 osmunda chandleri Last Modified in 12/03/21 | 8 | 39 | 37 | 0 | 0 | 0 | 0 | 3 |
| [105] † Osmunda nathorstii Last Modified in 12/03/21 | 27 | 18 | 18 | 0 | 0 | 0 | 0 | 3 |
| [106] † Osmunda oregonensis Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [107] † Osmunda pluma Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [108] † Osmunda precinnamomea Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [109] † Osmunda cinnamomea Neogene USA Last Modified in 12/03/21 | 3 | 42 | 42 | 0 | 0 | 0 | 0 | 3 |
| [110] † Osmunda cinnamomea Neogene Japan Last Modified in 12/03/21 | 6 | 39 | 39 | 0 | 0 | 0 | 0 | 3 |
| [111] † Osmunda iliaensis Last Modified in 12/03/21 | 1 | 45 | 43 | 0 | 0 | 0 | 0 | 3 |
| [112] † Osmunda shimokawaensis Last Modified in 12/03/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [113] † Osmunda wehrii Last Modified in 12/03/21 | 1 | 44 | 44 | 0 | 0 | 0 | 0 | 3 |
| [114] Osmunda cinnamomea Last Modified in 12/05/21 | 2 | 44 | 43 | 0 | 0 | 0 | 0 | 3 |
| [115] Osmunda claytoniana Last Modified in 12/05/21 | 1 | 46 | 44 | 0 | 0 | 0 | 0 | 3 |
| [116] Osmunda banksiifolia Last Modified in 12/05/21 | 2 | 44 | 42 | 0 | 0 | 0 | 0 | 3 |
| [117] Osmunda bromeliifolia Last Modified in 12/05/21 | 1 | 44 | 44 | 0 | 0 | 0 | 0 | 3 |
| [118] Osmunda javanica Last Modified in 12/05/21 | 1 | 46 | 43 | 0 | 0 | 0 | 0 | 3 |
| [119] Osmunda vachellii Last Modified in 12/05/21 | 2 | 43 | 43 | 0 | 0 | 0 | 0 | 3 |
| [120] Osmunda japonica Last Modified in 12/05/21 | 1 | 44 | 44 | 0 | 0 | 0 | 0 | 3 |
| [121] Osmunda lancea Last Modified in 12/05/21 | 1 | 45 | 44 | 0 | 0 | 0 | 0 | 3 |
| [122] Osmunda regalis Last Modified in 12/05/21 | 1 | 47 | 41 | 0 | 0 | 0 | 0 | 3 |
| [123] Leptopteris fraseri Last Modified in 12/05/21 | 2 | 45 | 41 | 0 | 0 | 0 | 0 | 3 |
| [124] Leptopteris hymenophylloides Last Modified in 12/05/21 | 2 | 44 | 43 | 0 | 0 | 0 | 0 | 3 |
| [125] Leptopteris wilkesiana Last Modified in 12/05/21 | 2 | 44 | 43 | 0 | 0 | 0 | 0 | 3 |
| [126] Leptopteris superba Last Modified in 12/05/21 | 2 | 44 | 43 | 0 | 0 | 0 | 0 | 3 |
| [127] Todea barbara Last Modified in 12/05/21 | 3 | 42 | 42 | 0 | 0 | 0 | 0 | 3 |
| [128] Todea papuana Last Modified in 12/05/21 | 38 | 7 | 7 | 0 | 0 | 0 | 0 | 3 |
| [129] † Osmundacaulis hoskingii tabulatus Last Modified in 12/03/21 | 7 | 38 | 37 | 0 | 0 | 0 | 0 | 3 |
| [130] † Millerocaulis australis Vera2010 Last Modified in 12/03/21 | 11 | 36 | 33 | 0 | 0 | 0 | 0 | 3 |
| [131] † Millerocaulis zamunerae Last Modified in 12/03/21 | 5 | 41 | 39 | 0 | 0 | 0 | 0 | 3 |
Project views
| type | number of views | Individual items viewed (where applicable) |
| Total project views | 6645 | |
| Project overview | 427 | |
| Media views | 1227 | Media search (1037 views); M834662 (190 views); |
| Matrix views | 356 | Matrix landing page (310 views); Morphological matrix from Bomfleur et al 2017 - (prepared by MorphoBank curators from original submission) (46 views); |
| Documents list | 323 | |
| Bibliography | 259 | |
| Taxon list | 2945 | |
| Specimen list | 684 | |
| Views for media list | 424 |
Project downloads
| type | number of downloads | Individual items downloaded (where applicable) |
| Total downloads from project | 15 | |
| Project downloads | 14 | |
| Matrix downloads | 1 | Morphological matrix from Bomfleur et al 2017 - (prepared by MorphoBank curators from original submission) (1 download); |