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Phylogenomic analyses in the complex Neotropical subfamily Corydoradinae (Siluriformes: Callichthyidae) with a new classification based on morphological and molecular data

Angelica C Dias, Luiz F C Tencatt, Fabio F Roxo, Gabriel de Souza da Costa Silva, Sérgio A Santos, Marcelo R Britto, Martin I Taylor, Claudio Oliveira
Zoological Journal of the Linnean Society, zlae053, https://doi.org/10.1093/zoolinnean/zlae053

Published: 11 June 2024


The family Callichthyidae, which comprises the subfamilies Corydoradinae and Callichthyinae, represents one of the largest families within the Siluriformes. Corydoradinae, the largest subfamily of Callichthyidae, alone accounts for >200 valid species, with new species being described frequently. This subfamily holds significant popularity among catfish enthusiasts worldwide, mainly because Corydoras are small, peaceful, and colourful fishes. Recognizing the existence of polyphyletic groups within Corydoradinae, the aim of this study was to construct a new phylogenetic hypothesis using genomic data (ultraconserved elements) and to re-evaluate the synonymized genera using the most recent morphological data. Our results supported the monophyly of Corydoradinae and identified seven distinct groups of species, each one with an associated pre-available generic name. These genera, namely Corydoras, Aspidoras, Scleromystax, Gastrodermus (resurrected), Osteogaster (resurrected), Brochis (resurrected), and Hoplisoma (resurrected), were all validated based on evidence derived from molecular and morphological analyses. A taxonomic key for the Corydoradinae genera is provided.

Classification breakdown supplied by "Fish in the news"

𝗟𝗶𝗻𝗲𝗮𝗴𝗲 𝗼𝗻𝗲 - which remains as 𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠.
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑐𝑢𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑚𝑎𝑝𝑎𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑟𝑒𝑖𝑜
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑢𝑟𝑜𝑓𝑒𝑛𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑙𝑜𝑐ℎ𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑎𝑟𝑎𝑚𝑎𝑡𝑒𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑒𝑟𝑣𝑖𝑛𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑟𝑖𝑎𝑡𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑟𝑡𝑒𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑒𝑠𝑎𝑛𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑓𝑙𝑎𝑚𝑒𝑛𝑡𝑜𝑠𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑓𝑜𝑤𝑙𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑓𝑢𝑙𝑙𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑒𝑜𝑓𝑓𝑟𝑜𝑦 (type species)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑎𝑐𝑢𝑙𝑖𝑓𝑒𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑎𝑟𝑐𝑖𝑠𝑠𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑒𝑔𝑟𝑜
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑢𝑟𝑎𝑠𝑡𝑖𝑔𝑚𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑥𝑦𝑟ℎ𝑦𝑛𝑐ℎ𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑠𝑡𝑎𝑧𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑎𝑟𝑎𝑚𝑎𝑐𝑐𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑎𝑟𝑎𝑟𝑒𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑒𝑚𝑖𝑎𝑞𝑢𝑖𝑙𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑒𝑝𝑡𝑒𝑛𝑡𝑟𝑖𝑜𝑛𝑎𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑒𝑟𝑟𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑖𝑚𝑢𝑙𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑜𝑙𝑜𝑥
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑝𝑖𝑙𝑢𝑟𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑡𝑒𝑛𝑜𝑐𝑒𝑝ℎ𝑎𝑙𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑡𝑟𝑒𝑖𝑡𝑙𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑣𝑖𝑡𝑎𝑡𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑧𝑎𝑤𝑎𝑑𝑧𝑘𝑖𝑖


𝗟𝗶𝗻𝗲𝗮𝗴𝗲 𝘁𝘄𝗼 - the 𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠.
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑎𝑙𝑏𝑎𝑡𝑒𝑟
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑎𝑙𝑑𝑒𝑏𝑎𝑟𝑎𝑛
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑎𝑧𝑎𝑔ℎ𝑎𝑙
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑏𝑒𝑙𝑒𝑛𝑜𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑏𝑟𝑢𝑛𝑛𝑒𝑢𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑐𝑎𝑟𝑣𝑎𝑙ℎ𝑜𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑑𝑒𝑝𝑖𝑛𝑛𝑎𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑓𝑢𝑠𝑐𝑜𝑔𝑢𝑡𝑎𝑡𝑢𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑔𝑎𝑏𝑟𝑖𝑒𝑙𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑘𝑖𝑟𝑖𝑟𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑘𝑜𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑚𝑎𝑐𝑢𝑙𝑜𝑠𝑢𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑚𝑒𝑝ℎ𝑖𝑠𝑡𝑜
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑝𝑜𝑒𝑐𝑖𝑙𝑢𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑝𝑠𝑎𝑚𝑚𝑎𝑡𝑖𝑑𝑒𝑠
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑟𝑎𝑖𝑚𝑢𝑛𝑑𝑖
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑟𝑜𝑐ℎ𝑎𝑖 (type species)
𝐴𝑠𝑝𝑖𝑑𝑜𝑟𝑎𝑠 𝑣𝑒𝑙𝑖𝑡𝑒𝑠


𝗟𝗶𝗻𝗲𝗮𝗴𝗲 𝘁𝗵𝗿𝗲𝗲 - the 𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥.
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑏𝑎𝑟𝑏𝑎𝑡𝑢𝑠 (type species)
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑙𝑎𝑐𝑒𝑟𝑑𝑎𝑖
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑚𝑎𝑐𝑟𝑜𝑝𝑡𝑒𝑟𝑢𝑠
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑝𝑟𝑖𝑜𝑛𝑜𝑡𝑜𝑠
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑟𝑒𝑖𝑠𝑖
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑠𝑎𝑙𝑚𝑎𝑐𝑖𝑠
𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑣𝑖𝑟𝑔𝑢𝑙𝑎𝑡𝑢𝑠


From the paper - "Our results corroborate the hypothesis of monophyly of 𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 proposed by Brito (2003) and Alexandrou et al. (2011), who named this assemblage lineage 3. All topologies demonstrate that 𝐶. 𝑙𝑎𝑐𝑒𝑟𝑑𝑎𝑖 belongs to this group and is the sister of all species in this genus. Specimens of 𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑐𝑒𝑟𝑑𝑎𝑖 were known only from its type series at the time of the studies by Reis (1998) and Brito (2003), and were not available in the original collections in the same period (Hieronimus 1995). Sampling efforts by some authors of the present paper (C.O. and M.R.B.) at the putative type locality resulted in a relatively large number of specimens that allowed examination of characters that recovered 𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑐𝑒𝑟𝑑𝑎𝑖 in 𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥, and herein we propose them combination 𝑆𝑐𝑙𝑒𝑟𝑜𝑚𝑦𝑠𝑡𝑎𝑥 𝑙𝑎𝑐𝑒𝑟𝑑𝑎𝑖. This combination appeared for the first time in the study by Fuller and Evers (2005: 272), although without any information to justify it."

𝗟𝗶𝗻𝗲𝗮𝗴𝗲𝘀 𝗳𝗼𝘂𝗿 𝗮𝗻𝗱 𝗳𝗶𝘃𝗲 are combined into 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠.
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑖𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑏𝑖𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑙𝑒𝑔𝑎𝑛𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑒𝑙𝑒𝑔𝑎𝑛𝑠 (type species)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑟𝑎𝑐𝑖𝑙𝑖𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑔𝑟𝑎𝑐𝑖𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑢𝑎𝑝𝑜𝑟𝑒 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑔𝑢𝑎𝑝𝑜𝑟𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 ℎ𝑎𝑠𝑡𝑎𝑡𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 ℎ𝑎𝑠𝑡𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑡𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑙𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑎𝑚𝑜𝑟𝑒 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑚𝑎𝑚𝑜𝑟𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑎𝑛𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑛𝑎𝑛𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑎𝑝𝑜𝑒𝑛𝑠𝑖𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑛𝑎𝑝𝑜𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑖𝑗𝑠𝑠𝑒𝑛𝑖 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑛𝑖𝑗𝑠𝑠𝑒𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑢𝑐𝑒𝑟𝑛𝑎 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑝𝑎𝑢𝑐𝑒𝑟𝑛𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑢𝑐𝑖𝑟𝑎𝑑𝑖𝑎𝑡𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑝𝑎𝑢𝑐𝑖𝑟𝑎𝑑𝑖𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑦𝑔𝑚𝑎𝑒𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑝𝑦𝑔𝑚𝑎𝑒𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑢𝑛𝑑𝑢𝑙𝑎𝑡𝑢𝑠 - 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 𝑢𝑛𝑑𝑢𝑙𝑎𝑡𝑢𝑠

From the paper - "𝑀𝑖𝑐𝑟𝑜𝑐𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 was proposed by Myers (1953) as a subgenus of 𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠, having 𝐺. ℎ𝑎𝑠𝑡𝑎𝑡𝑢𝑠 as its type species. Despite our results showing that 𝐺. ℎ𝑎𝑠𝑡𝑎𝑡𝑢𝑠 and 𝐺. 𝑝𝑦𝑔𝑚𝑎𝑒𝑢𝑠 form a clade sister group to the remaining 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠, we opted to keep 𝑀𝑖𝑐𝑟𝑜𝑐𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 in the synonymy of 𝐺𝑎𝑠𝑡𝑟𝑜𝑑𝑒𝑟𝑚𝑢𝑠 because the only putative diagnostic feature between these two genera would be the serration pattern on the posterior margin of the pectoral-fin spine."

𝗟𝗶𝗻𝗲𝗮𝗴𝗲 𝟳 are now 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟.
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑒𝑛𝑒𝑢𝑠 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑎𝑒𝑛𝑒𝑢𝑠 (type species)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑞𝑢𝑒𝑠 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑒𝑞𝑢𝑒𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 ℎ𝑒𝑝ℎ𝑎𝑒𝑠𝑡𝑢𝑠 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 ℎ𝑒𝑝ℎ𝑎𝑒𝑠𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑎𝑐𝑙𝑢𝑟𝑒𝑖 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑚𝑎𝑐𝑙𝑢𝑟𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑒𝑙𝑎𝑛𝑜𝑡𝑎𝑒𝑛𝑖𝑎 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑚𝑒𝑙𝑎𝑛𝑜𝑡𝑎𝑒𝑛𝑖𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑎𝑏𝑎𝑢𝑡𝑖 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑟𝑎𝑏𝑎𝑢𝑡𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑧𝑦𝑔𝑎𝑡𝑢𝑠 - 𝑂𝑠𝑡𝑒𝑜𝑔𝑎𝑠𝑡𝑒𝑟 𝑧𝑦𝑔𝑎𝑡𝑢𝑠


𝗟𝗶𝗻𝗲𝗮𝗴𝗲 𝟴 (comprising of subclades 1,2,3 & 4) are now 𝐵𝑟𝑜𝑐ℎ𝑖𝑠.
It is expected 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 will be revised in the future, leaving only Clade 1 species within i.e. 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠, etc.
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑔𝑎𝑠𝑠𝑖𝑧𝑖𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑎𝑔𝑎𝑠𝑠𝑖𝑧𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑚𝑎𝑛𝑑𝑎𝑗𝑎𝑛𝑒𝑎 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑎𝑚𝑎𝑛𝑑𝑎𝑗𝑎𝑛𝑒𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑚𝑏𝑖𝑎𝑐𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑎𝑚𝑏𝑖𝑎𝑐𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑝𝑝𝑟𝑜𝑢𝑎𝑔𝑢𝑒𝑛𝑠𝑖𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑎𝑝𝑝𝑟𝑜𝑢𝑎𝑔𝑢𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑟𝑐𝑢𝑎𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑎𝑟𝑐𝑢𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑒𝑡ℎ𝑎𝑛𝑎𝑒 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑏𝑒𝑡ℎ𝑎𝑛𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑖𝑓𝑎𝑠𝑐𝑖𝑎𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑏𝑖𝑓𝑎𝑠𝑐𝑖𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑟𝑖𝑡𝑠𝑘𝑖𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑏𝑟𝑖𝑡𝑠𝑘𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑟𝑖𝑡𝑜𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑏𝑟𝑖𝑡𝑜𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑛𝑑𝑖𝑠𝑐𝑖𝑝𝑢𝑙𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑐𝑜𝑛𝑑𝑖𝑠𝑐𝑖𝑝𝑢𝑙𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑠𝑡𝑎𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑐𝑜𝑠𝑡𝑎𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑟𝑖𝑚𝑚𝑒𝑛𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑐𝑟𝑖𝑚𝑚𝑒𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑟𝑦𝑝𝑡𝑖𝑐𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑐𝑟𝑦𝑝𝑡𝑖𝑐𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑒𝑙𝑝ℎ𝑎𝑥 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑑𝑒𝑙𝑝ℎ𝑎𝑥
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑒𝑤𝑒𝑦𝑒𝑟𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑑𝑒𝑤𝑒𝑦𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑖𝑓𝑢𝑣𝑖𝑎𝑡𝑖𝑙𝑖𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑑𝑖𝑓𝑢𝑣𝑖𝑎𝑡𝑖𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑝ℎ𝑖𝑝𝑝𝑖𝑓𝑒𝑟 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑒𝑝ℎ𝑖𝑝𝑝𝑖𝑓𝑒𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑎𝑟𝑏𝑒𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑔𝑎𝑟𝑏𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑒𝑟𝑦𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑔𝑒𝑟𝑦𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑜𝑚𝑒𝑧𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑔𝑜𝑚𝑒𝑧𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 ℎ𝑎𝑟𝑎𝑙𝑑𝑠𝑐ℎ𝑢𝑙𝑡𝑧𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 ℎ𝑎𝑟𝑎𝑙𝑑𝑠𝑐ℎ𝑢𝑙𝑡𝑧𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 ℎ𝑒𝑡𝑒𝑟𝑜𝑚𝑜𝑟𝑝ℎ𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 ℎ𝑒𝑡𝑒𝑟𝑜𝑚𝑜𝑟𝑝ℎ𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑖𝑚𝑖𝑡𝑎𝑡𝑜𝑟 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑖𝑚𝑖𝑡𝑎𝑡𝑜𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑖𝑛𝑐𝑜𝑙𝑖𝑐𝑎𝑛𝑎 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑖𝑛𝑐𝑜𝑙𝑖𝑐𝑎𝑛𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑖𝑠𝑏𝑟𝑢𝑒𝑐𝑘𝑒𝑟𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑖𝑠𝑏𝑟𝑢𝑒𝑐𝑘𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑚𝑏𝑒𝑟𝑡𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑙𝑎𝑚𝑏𝑒𝑟𝑡𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑒𝑜𝑝𝑎𝑟𝑑𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑙𝑒𝑜𝑝𝑎𝑟𝑑𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑢𝑙𝑡𝑖𝑟𝑎𝑑𝑖𝑎𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑚𝑢𝑙𝑡𝑖𝑟𝑎𝑑𝑖𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑜𝑒𝑙𝑘𝑒𝑚𝑝𝑓𝑓𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑛𝑜𝑒𝑙𝑘𝑒𝑚𝑝𝑓𝑓𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑟𝑛𝑎𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑜𝑟𝑛𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑟𝑝ℎ𝑛𝑜𝑝𝑡𝑒𝑟𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑜𝑟𝑝ℎ𝑛𝑜𝑝𝑡𝑒𝑟𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑛𝑡𝑎𝑛𝑎𝑙𝑒𝑛𝑠𝑖𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑝𝑎𝑛𝑡𝑎𝑛𝑎𝑙𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑖𝑛ℎ𝑒𝑖𝑟𝑜𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑝𝑖𝑛ℎ𝑒𝑖𝑟𝑜𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑢𝑙𝑐ℎ𝑒𝑟 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑝𝑢𝑙𝑐ℎ𝑒𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑒𝑡𝑖𝑐𝑢𝑙𝑎𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑟𝑒𝑡𝑖𝑐𝑢𝑙𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑜𝑏𝑖𝑛𝑒𝑎𝑒 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑟𝑜𝑏𝑖𝑛𝑒𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑜𝑏𝑢𝑠𝑡𝑢𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑟𝑜𝑏𝑢𝑠𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑒𝑢𝑠𝑠𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑒𝑢𝑠𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑜𝑑𝑎𝑙𝑖𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑜𝑑𝑎𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑝𝑒𝑐𝑡𝑎𝑏𝑖𝑙𝑖𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑝𝑒𝑐𝑡𝑎𝑏𝑖𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠 (type species)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑦𝑐ℎ𝑟𝑖 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑦𝑐ℎ𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑣𝑖𝑟𝑔𝑖𝑛𝑖𝑎𝑒 - 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑣𝑖𝑟𝑔𝑖𝑛𝑖𝑎𝑒

From the paper - "Our results revealed the putative existence of two species of 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠 (Castelnau, 1855): one from Peru (𝐵. 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠, voucher 87580, LBP 22528) and other from Paraguay River Basin (𝐵. 𝑎𝑓𝑓. 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠, voucher 41751, LBP 8503). 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠 is described originally from Tocantins River, and there is an available name for species from Peru currently synonymized to 𝐵. 𝑠𝑝𝑙𝑒𝑛𝑑𝑒𝑛𝑠, 𝐵𝑟𝑜𝑐ℎ𝑖𝑠 𝑐𝑜𝑒𝑟𝑢𝑙𝑒𝑢𝑠 (Cope, 1872), described from Ampiyacu river, the same river where the sample that we have in our analysis was collected."

𝗟𝗶𝗻𝗲𝗮𝗴𝗲𝘀 𝟲 𝗮𝗻𝗱 𝟵 are now 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎.
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑐𝑟𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑐𝑟𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑑𝑜𝑙𝑓𝑜𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑑𝑜𝑙𝑓𝑜𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑙𝑏𝑜𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑙𝑏𝑜𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑚𝑝ℎ𝑖𝑏𝑒𝑙𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑚𝑝ℎ𝑖𝑏𝑒𝑙𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑝𝑖𝑎𝑘𝑎 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑝𝑖𝑎𝑘𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑟𝑎𝑔𝑢𝑎𝑖𝑎𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑟𝑎𝑔𝑢𝑎𝑖𝑎𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑟𝑚𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑟𝑚𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑡𝑟𝑜𝑝𝑒𝑟𝑠𝑜𝑛𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑡𝑟𝑜𝑝𝑒𝑟𝑠𝑜𝑛𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑎𝑥𝑒𝑙𝑟𝑜𝑑𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑎𝑥𝑒𝑙𝑟𝑜𝑑𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑎𝑑𝑒𝑟𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑎𝑑𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑒𝑛𝑎𝑡𝑖𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑒𝑛𝑎𝑡𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑖𝑐𝑜𝑙𝑜𝑟 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑖𝑐𝑜𝑙𝑜𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑜𝑒ℎ𝑙𝑘𝑒𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑜𝑒ℎ𝑙𝑘𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑜𝑒𝑠𝑒𝑚𝑎𝑛𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑜𝑒𝑠𝑒𝑚𝑎𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑜𝑛𝑑𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑜𝑛𝑑𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑟𝑒𝑒𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑟𝑒𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑟𝑒𝑣𝑖𝑟𝑜𝑠𝑡𝑟𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑟𝑒𝑣𝑖𝑟𝑜𝑠𝑡𝑟𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑏𝑢𝑟𝑔𝑒𝑠𝑠𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑏𝑢𝑟𝑔𝑒𝑠𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑎𝑟𝑙𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑎𝑟𝑙𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑎𝑢𝑑𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑎𝑢𝑑𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑐ℎ𝑢𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑜𝑐ℎ𝑢𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑙𝑜𝑠𝑠𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑜𝑙𝑜𝑠𝑠𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑛𝑐𝑜𝑙𝑜𝑟 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑜𝑛𝑐𝑜𝑙𝑜𝑟
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑝𝑒𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑜𝑝𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑜𝑝𝑝𝑒𝑛𝑎𝑚𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑜𝑝𝑝𝑒𝑛𝑎𝑚𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑐𝑟𝑢𝑧𝑖𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑐𝑟𝑢𝑧𝑖𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑎𝑣𝑖𝑑𝑠𝑎𝑛𝑑𝑠𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑑𝑎𝑣𝑖𝑑𝑠𝑎𝑛𝑑𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑖𝑝ℎ𝑦𝑒𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑑𝑖𝑝ℎ𝑦𝑒𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑑𝑢𝑝𝑙𝑖𝑐𝑎𝑟𝑒𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑑𝑢𝑝𝑙𝑖𝑐𝑎𝑟𝑒𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒ℎ𝑟ℎ𝑎𝑟𝑑𝑡𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑒ℎ𝑟ℎ𝑎𝑟𝑑𝑡𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑠𝑝𝑒𝑟𝑎𝑛𝑧𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑒𝑠𝑝𝑒𝑟𝑎𝑛𝑧𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑣𝑒𝑙𝑦𝑛𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑒𝑣𝑒𝑙𝑦𝑛𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑒𝑣𝑒𝑟𝑠𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑒𝑣𝑒𝑟𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑓𝑎𝑣𝑒𝑜𝑙𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑓𝑎𝑣𝑒𝑜𝑙𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑓𝑜𝑒ℎ𝑙𝑖𝑐ℎ𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑓𝑜𝑒ℎ𝑙𝑖𝑐ℎ𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑙𝑎𝑑𝑦𝑠𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑙𝑎𝑑𝑦𝑠𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑜𝑠𝑠𝑒𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑜𝑠𝑠𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑟𝑎𝑛𝑡𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑟𝑎𝑛𝑡𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑟𝑖𝑠𝑒𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑟𝑖𝑠𝑒𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑟𝑦𝑝ℎ𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑟𝑦𝑝ℎ𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑔𝑢𝑖𝑎𝑛𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑔𝑢𝑖𝑎𝑛𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 ℎ𝑎𝑏𝑟𝑜𝑠𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 ℎ𝑎𝑏𝑟𝑜𝑠𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑗𝑢𝑙𝑖𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑗𝑢𝑙𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑘𝑎𝑛𝑒𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑘𝑎𝑛𝑒𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑘𝑛𝑎𝑎𝑐𝑘𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑘𝑛𝑎𝑎𝑐𝑘𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑎𝑐𝑟𝑖𝑚𝑜𝑠𝑡𝑖𝑔𝑚𝑎𝑡𝑎 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑎𝑐𝑟𝑖𝑚𝑜𝑠𝑡𝑖𝑔𝑚𝑎𝑡𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑒𝑢𝑐𝑜𝑚𝑒𝑙𝑎𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑒𝑢𝑐𝑜𝑚𝑒𝑙𝑎𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑜𝑛𝑔𝑖𝑝𝑖𝑛𝑛𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑜𝑛𝑔𝑖𝑝𝑖𝑛𝑛𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑜𝑟𝑒𝑡𝑜𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑜𝑟𝑒𝑡𝑜𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑜𝑥𝑜𝑧𝑜𝑛𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑜𝑥𝑜𝑧𝑜𝑛𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑙𝑦𝑚𝑛𝑎𝑑𝑒𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑙𝑦𝑚𝑛𝑎𝑑𝑒𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑒𝑙𝑎𝑛𝑖𝑠𝑡𝑖𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑒𝑙𝑎𝑛𝑖𝑠𝑡𝑖𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑒𝑙𝑖𝑛𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑒𝑙𝑖𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑒𝑡𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑒𝑡𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑖𝑐𝑟𝑎𝑐𝑎𝑛𝑡ℎ𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑖𝑐𝑟𝑎𝑐𝑎𝑛𝑡ℎ𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑖𝑐𝑟𝑜𝑐𝑒𝑝ℎ𝑎𝑙𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑖𝑐𝑟𝑜𝑐𝑒𝑝ℎ𝑎𝑙𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑚𝑢𝑙𝑡𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑚𝑢𝑙𝑡𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑛𝑎𝑡𝑡𝑒𝑟𝑒𝑟𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑛𝑎𝑡𝑡𝑒𝑟𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑖𝑎𝑝𝑜𝑞𝑢𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑜𝑖𝑎𝑝𝑜𝑞𝑢𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑟𝑡𝑒𝑔𝑎𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑜𝑟𝑡𝑒𝑔𝑎𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑜𝑠𝑡𝑒𝑜𝑐𝑎𝑟𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑜𝑠𝑡𝑒𝑜𝑐𝑎𝑟𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑙𝑒𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑎𝑙𝑒𝑎𝑡𝑢𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑛𝑑𝑎 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑎𝑛𝑑𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑟𝑎𝑔𝑢𝑎 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑎𝑟𝑎𝑔𝑢𝑎
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑟𝑎𝑙𝑙𝑒𝑙𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑎𝑟𝑎𝑙𝑙𝑒𝑙𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑎𝑣𝑎𝑛𝑒𝑙𝑙𝑖𝑎𝑒 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑎𝑣𝑎𝑛𝑒𝑙𝑙𝑖𝑎𝑒
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑒𝑡𝑟𝑎𝑐𝑖𝑛𝑖𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑒𝑡𝑟𝑎𝑐𝑖𝑛𝑖𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑜𝑙𝑦𝑠𝑡𝑖𝑐𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑜𝑙𝑦𝑠𝑡𝑖𝑐𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑜𝑡𝑎𝑟𝑜𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑜𝑡𝑎𝑟𝑜𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑝𝑢𝑛𝑐𝑡𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑝𝑢𝑛𝑐𝑡𝑎𝑡𝑢𝑚 (type species)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑒𝑣𝑒𝑙𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑟𝑒𝑣𝑒𝑙𝑎𝑡𝑢𝑚 (fossil record)
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑟𝑒𝑦𝑛𝑜𝑙𝑑𝑠𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑟𝑒𝑦𝑛𝑜𝑙𝑑𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑎𝑛𝑐ℎ𝑒𝑠𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑎𝑛𝑐ℎ𝑒𝑠𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑐ℎ𝑤𝑎𝑟𝑡𝑧𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑐ℎ𝑤𝑎𝑟𝑡𝑧𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑖𝑚𝑖𝑙𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑖𝑚𝑖𝑙𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑖𝑝𝑎𝑙𝑖𝑤𝑖𝑛𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑖𝑝𝑎𝑙𝑖𝑤𝑖𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑡𝑒𝑖𝑛𝑑𝑎𝑐ℎ𝑛𝑒𝑟𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑡𝑒𝑖𝑛𝑑𝑎𝑐ℎ𝑛𝑒𝑟𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑡𝑒𝑟𝑏𝑎𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑡𝑒𝑟𝑏𝑎𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑠𝑢𝑟𝑖𝑛𝑎𝑚𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑠𝑢𝑟𝑖𝑛𝑎𝑚𝑒𝑛𝑠𝑖𝑠
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑡𝑟𝑖𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑡𝑟𝑖𝑙𝑖𝑛𝑒𝑎𝑡𝑢𝑚
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑡𝑢𝑘𝑎𝑛𝑜 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑡𝑢𝑘𝑎𝑛𝑜
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑢𝑟𝑢𝑐𝑢 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑢𝑟𝑢𝑐𝑢
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑤𝑒𝑖𝑡𝑧𝑚𝑎𝑛𝑖 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑤𝑒𝑖𝑡𝑧𝑚𝑎𝑛𝑖
𝐶𝑜𝑟𝑦𝑑𝑜𝑟𝑎𝑠 𝑥𝑖𝑛𝑔𝑢𝑒𝑛𝑠𝑖𝑠 - 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎 𝑥𝑖𝑛𝑔𝑢𝑒𝑛𝑠𝑖𝑠


Note some species names, the authors had to change as adjectives, i.e. 𝑐𝑎𝑢𝑑𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑠 becomes 𝑐𝑎𝑢𝑑𝑖𝑚𝑎𝑐𝑢𝑙𝑎𝑡𝑢𝑚. Only applied in 𝐻𝑜𝑝𝑙𝑖𝑠𝑜𝑚𝑎.







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