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Redescription of the Far Eastern Brook Lamprey Lethenteron Reissneri (Dybowski, 1869) (Petromyzontidae)

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Redescription of the Far Eastern Brook Lamprey Lethenteron Reissneri (Dybowski, 1869) (Petromyzontidae) A peer-reviewed open-access journal ZooKeys 506: 75–93Redescription (2015) of the Far Eastern brook lamprey Lethenteron reissneri... 75 doi: 10.3897/zookeys.506.9817 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research Redescription of the Far Eastern brook lamprey Lethenteron reissneri (Dybowski, 1869) (Petromyzontidae) Claude B. Renaud1, Alexander M. Naseka2 1 Research & Collections Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, ON K1P 6P4 Canada 2 Faculty for Biology and Soil, St. Petersburg State University, Universitetskaya Emb. 7, St. Petersburg, 199034 Russia Corresponding author: Claude B. Renaud ([email protected]) Academic editor: N. Bogutskaya | Received 17 April 2015 | Accepted 18 May 2015 | Published 28 May 2015 http://zoobank.org/6055196F-5DF5-4FF1-8C2D-1740E4DC70BC Citation: Renaud CB, Naseka AM (2015) Redescription of the Far Eastern brook lamprey Lethenteron reissneri (Dybowski, 1869) (Petromyzontidae). ZooKeys 506: 75–93. doi: 10.3897/zookeys.506.9817 Abstract Nonparasitic Lethenteron reissneri (Dybowski) is redescribed based on four syntypes (two adults and two ammocoetes) from the Onon and Ingoda rivers, Russia, and 15 topotypic specimens (seven metamor- phosing ammocoetes and eight ammocoetes) from the Onon River system, Russia and Mongolia. Poste- rial teeth were not mentioned in the original description, but Berg (1931) stated that they were sometimes absent, which he later (Berg 1948) changed to usually absent, based on material (some of which we have re-identified as parasitic L. camtschaticum) from far outside of the type locality. The latter view has been widely accepted by subsequent authors. Unfortunately, the poor condition of the two adult syntypes did not permit verification of this character. However, a row of posterials was clearly visible in six of the seven topotypic metamorphosing ammocoetes and indicates their usual presence in the species. The first full description of the ammocoetes, including pigmentation, is provided. The present study restricts the distri- bution of L. reissneri to the Shilka and Songhua river systems within the Amur River basin, until a more geographically comprehensive study is undertaken. Additionally, in this study, feeding versus non-feeding at the adult stage, are considered to be valid taxonomic characters at the species level. Keywords Amur River basin, morphology, nonparasitic, taxonomy Copyright Claude B. Renaud, Alexander M. Naseka. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 76 Claude B. Renaud & Alexander M. Naseka / ZooKeys 506: 75–93 (2015) Introduction The Northern Hemisphere lamprey genus Lethenteron was originally erected as a sub- genus of Entosphenus by Creaser and Hubbs (1922) and elevated to generic rank by Jordan et al. (1930), without justification. Vladykov and Follett (1967) accepted this action and defined members of Lethenteron as possessing posterial teeth in a single curved row and with the outer laterals (exolaterals) absent. Renaud (2011) recognized one parasitic species, L. camtschaticum (Tilesius, 1811) (i.e., the stem species) and six nonparasitic species, L. alaskense Vladykov & Kott, 1978, L. appendix (DeKay, 1842), L. kessleri (Anikin, 1905), L. ninae Naseka, Tuniyev & Renaud, 2009, L. reissneri (Dybowski, 1869), and L. zanandreai (Vladykov, 1955) (i.e., the satellite species) in the genus, along with two additional undescribed nonparasitic species from Japan, and noted that the taxonomic limits of the nonparasitic L. reissneri were unclear. Authors have either treated these satellites of L. camtschaticum as distinct species (e.g., Potter et al. 2015) or as synonyms of this species. For example, Artamonova et al. (2011a, 2011b) and Makhrov et al. (2013) have, respectively, suggested that L. ninae, L. re- issneri, and L. kessleri are synonyms of L. camtschaticum and that mode of life (i.e., parasitism versus nonparasitism) is not a valid criterion for specific distinctiveness. This is not a new concept. McPhail and Lindsey (1970), in discussing L. camtschaticum (reported as Lampetra japonica (von Martens, 1868)) from the Northwest Territories, Yukon and Alaska, suggested that it may not be specifically distinct fromL . appendix (reported as L. lamottenii (Lesueur, 1827)) from eastern North America, but preferred to retain them as distinct pending a review of all relevant species from both contin- ents (i.e., North America and Eurasia). Renaud et al. (2009) proposed that the key to resolving the issue of whether parasitism or nonparasitism constitute valid taxonomic characters at the species level may be to conduct common garden experiments (i.e., rearing two putative species under the same conditions from the zygote to the adult stage) that seek to elucidate the triggering mechanism for trophic interactions, or the lack thereof, in the adult stage. In the absence of such experiments, however, we follow here conventional taxonomy and continue to recognize the above nonparasitic species as distinct. Additionally, prior to synonymizing species, it is critical to compare any extant type material as well as the original descriptions, and, unfortunately, this has not been done on a consistent basis by authors. The original description by Dybowski (1869) of Petromyzon reissneri was very short, incomplete and based on material from the upper Amur River basin, Russia (type locality: Onon and Ingoda rivers). A number of authors conducted morphological studies on what they referred to as Lampetra reiss- neri and anadromous L. japonica (= Lethenteron camtschaticum) from the lower Amur River basin (Berg 1931, 1948, Nikol’sky 1956, Abakumov 1960, Hensel 1963). How- ever, we are aware of only two studies since the original description, to have examined lamprey material, which they treated as a single species, from the upper Amur (Karasev 1987, Yamazaki et al. 2006). The Amur River basin is divided into five or six well- defined zoogeographic districts (Bogutskaya et al. 2008) or ecoregions (Abell et al. Redescription of the Far Eastern brook lamprey Lethenteron reissneri... 77 2008), respectively, based on the distributional disparities of freshwater fish species along its 4,370–4,510 km length. Thus, it is critical to examine lamprey material from the upper rather than lower Amur, to objectively evaluate the characteristics of L. reiss- neri and prevent the inclusion of another species in the redescription. Karasev (1987) studied 52 adult lamprey from the Ingoda and the Shilka rivers, but only reported a few morphometric characters and the tooth formula of the infraoral lamina. Yamazaki et al. (2006) studied 26 ammocoetes from the Onon and 29 from the Ingoda rivers and determined that they possessed 65–73 trunk myomeres, but made no mention of their body pigmentation. Yamazaki et al. (2006) examined an adult syntype of L. re- issneri (Museum für Naturkunde, Berlin; ZMB 7118) and determined it had 69 trunk myomeres, but made no mention of its dentition. Although Berg (1931) reported that L. reissneri possessed 56–67 trunk myomeres, that range was taken from Jordan and Hubbs (1925) based on Japanese material (Himeji and Lake Biwa, Honshu Island and Sapporo, Hokkaido Island), which they identified as Entosphenus mitsukurii (Hatta 1901). Berg (1931) further stated that the lower labial teeth (= posterial teeth) were sometimes absent in L. reissneri. He subsequently (Berg 1948) changed this to usu- ally absent based on material from the Asian Pacific coast between the Anadyr estu- ary and the Shangshi River, distant from the type locality. Hubbs and Potter (1971) highlighted the inconsistency between Berg’s (1948) statement of usually no lower labial teeth and their observations. Nevertheless, Reshetnikov (2002) repeated Berg’s (1948) statement in the widely cited Atlas of freshwater fishes of Russia. The purpose of this study is to provide a full description of the species based on the examination of type and topotypic material of ammocoetes, metamorphosing ammocoetes and adults, and in view of the inconsistency between the observations of Berg (1931, 1948) and Hubbs and Potter (1971), re-examine material that Berg studied to determine how often posterials are absent. Materials and methods Material was examined from the Academy of Natural Sciences, Philadelphia (ANSP), Natural History Museum, London (BMNH), Canadian Museum of Nature Fish Col- lection, Ottawa (CMNFI), Naturhistorisches Museum Wien, Vienna (NMW), Zoo- logical Institute, Russian Academy of Sciences, St. Petersburg (ZIN), and Museum für Naturkunde, Berlin (ZMB). Note that collection dates for ZIN collections use the Gregorian rather than the Julian calendar. The characters examined in the ammocoetes and adults follow the method of Renaud (2011). Evaluation of the lateral line neuro- mast pigmentation in both ammocoetes and adults was made on the dorsal aspect of the branchial region. We additionally examined in ammocoetes the pigment pattern of the posterior fleshy part of the tail as in Richards et al. (1982); their caudal ridge, and the pigmentation in adults of the gular region and apex of the second dorsal fin following Vladykov and Kott (1978). TL, total length. 78 Claude B. Renaud & Alexander M. Naseka / ZooKeys 506: 75–93 (2015) Figure 1. Geographic distribution of Lethenteron reissneri and material identified
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