Updated Morphological Descriptions of the Larval Stage of Urophycis (Family: Phycidae) from the Northeast United States Continental Shelf
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Updated Morphological Descriptions of the Larval Stage of Urophycis (Family: Phycidae) from the Northeast United States Continental Shelf Authors: Marancik, Katrin E., Richardson, David E., and Konieczna, Małgorzata Source: Copeia, 108(1) : 83-90 Published By: The American Society of Ichthyologists and Herpetologists URL: https://doi.org/10.1643/CG-19-219 BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Complete website, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/terms-of-use. Usage of BioOne Complete content is strictly limited to personal, educational, and non - commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Downloaded From: https://bioone.org/journals/Copeia on 13 Aug 2021 Terms of Use: https://bioone.org/terms-of-use Copeia 108, No. 1, 2020, 83–90 Updated Morphological Descriptions of the Larval Stage of Urophycis (Family: Phycidae) from the Northeast United States Continental Shelf Katrin E. Marancik1, David E. Richardson2, and Małgorzata Konieczna3 Including early life history data in assessments can improve fisheries management by increasing our knowledge of stock structure, spawning habitat, and population trends. The identification of fish larvae to species is a necessary step in using early life history data toward this goal. Three species of hakes from the genus Urophycis are common on the northeast United States continental shelf: U. chuss or Red Hake, U. regia or Spotted Hake, and U. tenuis or White Hake. Unfortunately, identification of larval Urophycis has long been only possible at the genus level. Larvae of Urophycis (n ¼ 277) collected in a subset of ethanol-preserved samples were identified genetically through sequencing of the cytochrome oxidase I gene and were used to update morphological descriptions with characters that separate these three species at the larval stage. Sequencing occurred in two stages: the first (n ¼ 88) to develop a set of known-identity larvae to define species-specific traits, the second (n ¼ 189) to test morphological identification based on the traits described in this study. We describe a combination of the location of dorsal and ventral pigment, head pigment, lower jaw pigment, and the timing of development of the pectoral fins to distinguish the larvae of these three species at sizes ,6 mm. Using molecular techniques to improve morphological identifications is a powerful and efficient way to obtain the species-level data needed for assessments and management. IFFICULTY identifying larval-stage hakes of the examine those larvae for species-specific traits, and then genus Urophycis (U. chuss, U. regia, and U. tenuis)to to test the accuracy of identifications made with those D species has limited the use of over 40 years of traits. fisheries-independent data collected on these taxa on the northeast United States continental shelf. Larvae of Urophycis MATERIALS AND METHODS are among the most abundant ichthyoplankton taxa collect- ed in this region (Walsh et al., 2015). The frequency and long Sincethelate1970s,fishlarvaehavebeencollected duration of plankton monitoring on the northeast U.S. shelf through the Marine Resources Monitoring, Assessment, means these data could provide important insights into the and Prediction (MARMAP) and Ecosystem Monitoring population structure of these species and an independent (EcoMon) programs of the National Oceanic and Atmo- estimate of trends in spawning stock biomass (Richardson et spheric Administration’s (NOAA) Northeast Fisheries Sci- al., 2009). ence Center, which survey plankton, hydrography, and Larvae of Urophycis are morphologically similar, and water chemistry on the entire continental shelf from Cape previous descriptions did not produce reliable identifying Hatteras, North Carolina, to Cape Sable, Nova Scotia. The characteristics at the species level for small larvae. shelf is divided into strata based on bathymetry and Individual species have been described in detail based on latitude and sampled using a random stratified sampling reared larvae from known adults (U. chuss: Miller and design. For each survey, stations are randomly selected Marak, 1959; U. regia: Barans and Barans, 1972 and within strata with up to six surveys per year: January/ Serebryakov, 1978), but comparisons among the common February, March–April, May/June, August, September–Oc- species relied heavily on characters that are unavailable at tober, and November. Samples are collected with oblique the small sizes common in plankton tows (,6 mm) or tows of a 61 cm diameter bongo net with 505 lm characters that could be misleading (i.e., the presence of (MARMAP) or 333 lm (EcoMon) mesh, which is towed to pelvic filament pigment is unreliable; Methven, 1985; within 5 m of the bottom or to a maximum depth of 200 Comyns and Grant, 1993; Comyns and Bond, 2002; Fahay, m (Walsh et al., 2015). All standard samples are preserved 2007). in 5% buffered formalin and are sorted and initially Genetic barcoding provides a means to identify larvae at identified at the Plankton Sorting and Identification the species level (e.g., Marancik et al., 2010). Although Center (ZSIOP) in Szczecin, Poland. Since 2013, additional quite effective, genetic identification of entire surveys samples have been collected at each station during would be costly. However, genetic barcoding of a subset EcoMon surveys with a 20 cm diameter bongo with 333 of specimens is a relatively inexpensive way to improve lm mesh nets attached above the 61 cm bongo frame. and test the accuracy of identifications made through These additional samples were preserved in 95% ethanol, morphological analysis. sorted and identified in house, and used for genetic TheobjectivesofthisstudyaretouseDNAbarcodingto identification of eggs (Lewis et al., 2016) and larvae of create a set of larvae of Urophycis of known identity, to species of Urophycis. 1 Integrated Statistics, 28 Tarzwell Drive, Narragansett, Rhode Island 02882; Email: [email protected]. Send reprint requests to this address. 2 NOAA Fisheries Northeast Fisheries Science Center, 28 Tarzwell Drive, Narragansett, Rhode Island 02882; Email: [email protected]. 3 Morski Instytut Rybacki, Plankton Sorting and Identification Center (ZSIOP), Szczecin, Poland. Submitted: 2 April 2019. Accepted: 4 December 2019. Associate Editor: M. P. Davis. Ó 2020 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CG-19-219 Published online: 20 February 2020 Downloaded From: https://bioone.org/journals/Copeia on 13 Aug 2021 Terms of Use: https://bioone.org/terms-of-use 84 Copeia 108, No. 1, 2020 In addition to the EcoMon samples, a cruise in May 1993 Table 1. Confusion matrix comparing morphological identifications to on the NOAA ship Albatross IV sampled off the U.S. mid- molecular results for the two rounds of barcoding. Failed refers to No Atlantic shelf edge, the suspected spawning ground for sequence or Low-Quality sequences amplified during multiple rounds Urophycis tenuis (Fahay and Able, 1989; Lang et al., 1996). of barcoding. Round 1 included 88 larvae morphologically identified to genus and used to define species-level traits. Round 2 included 189 Larvae collected in these off-shelf samples had morpho- larvae morphologically identified to species and used to test identifi- logical differences from the shelf larvae collected in cations based on the traits described in this study. August–November, so they are suspected to be U. tenuis. Unfortunately, these samples were preserved in 5% Molecular results Morphological buffered formalin and were not available for genetic results U. chuss U. regia U. tenuis Failed Total barcoding. Therefore, these specimens were insufficient for unequivocal identification. Round 1 For genetic barcoding, eyeballs from ethanol-preserved Urophycis spp. 80 6 0 2 88 larvae were removed in the lab, placed in a 96-well plate, and Round 2 shipped to the University of Guelph Canadian Centre for U. chuss 108 1 0 17 126 DNA Barcoding. Genetic identification procedures were U. regia 1430 549 based on the Barcode of Life protocols amplifying a 650 bp U. tenuis 000 00 section and a 184 bp section of the cytochrome oxidase I Urophycis spp. 7 3 0 4 14 gene (Ivanova et al., 2012; Lewis et al., 2016). Total 196 53 0 28 277 Larvae were barcoded in two groups. The first group (n ¼ 88) were morphologically identified to genus. After barcoding, these larvae of known identity were used to define species-specific traits for morphological identifica- collections of Urophycis) based on examination of ethanol- tion. The second group of larvae (n ¼ 189) were and formalin-preserved larvae. While pigment was present in morphologically identified to species, and then these other regions (e.g., nape, gut), the individual-level variability identifications were tested with barcoding to confirm was sufficient to preclude this from being useful for species- validity of the traits described for separating species.