Morphological and genetic structuring in the Utah Lake sucker complex

Population decline in the federally endangered June sucker (Chasmistes liorus), a lakesucker unique to Utah Lake, Utah, has been attributed in part to hybridization with the more widespread Utah sucker (Catostomus ardens). As a group, suckers in Utah Lake exhibit considerable external morphological variation. Meristic and morphological ambiguities, presumably the result of hybridization, create a continuum of intermediate forms between Chasmistes and Catostomus extremes and prevent definitive identification to species. Here we describe and evaluate the morphological and genetic variation in suckers in Utah Lake by comparing a morphological analysis with amplified fragment length polymorphism and microsatellite analyses. Suckers were morphologically differentiated using mouth characters associated with different feeding strategies: planktivory (June sucker) and benthivory (Utah sucker). Although we found no genetic evidence for a deep divergence between June and Utah morphs, significant, but slight population structuring accompanied the substantial morphological variation. Bayesian model‐based genetic clustering analyses detected two sucker populations in Utah Lake; however, these clusters were not strongly concordant with morphological groupings or between marker systems. The suckers in Utah Lake present an interesting dilemma regarding conservation: should one conserve (breed and stock) a subset of the morphotypic variation in the Utah Lake sucker complex, focusing on the endangered June sucker morphotype, or should one conserve both June sucker and Utah sucker morphotypes in this complex, possibly maximizing evolutionary potential? We explore this question in the context of current genetic and morphological variation in the Utah Lake sucker complex as well as historical information on this complex and other lakesuckers.     

Evolution after the flood: phylogeography of the desert fish Utah Chub

The Bonneville Basin and upper Snake River drainage of western North America underwent extensive hydrological changes during the late Pleistocene, potentially influencing the geographic distribution and evolutionary trajectories of aquatic species that occupied this region. To test this hypothesis, I reconstructed the phylogeographic history of the desert fish Utah chub (Gila atraria) by examining 16 populations that span the natural distribution of this species across the Bonneville Basin and upper Snake River. I compared mitochondrial control region sequences (934 bp) among 77 individuals revealing 24 unique haplotypes. Geographic and phylogenetic relationships among haplotypes were explored using parsimony, maximum likelihood, nested clade analysis, and analysis of molecular variance. I found that G. atraria is composed of two distinct clades that represent an early Pleistocene split between the upper Snake River and Bonneville Basin. Within each of these clades, geographic structuring was highly concordant with the hydrological history of late Pleistocene Lake Bonneville and the upper Snake River, suggesting that glacial-induced shifts in climate and unpredictable geological events have played a major role in shaping genetic subdivision among populations. To examine the effects of vicariant events on phenotypic divergence among Utah chub populations, I mapped chub life histories to the control region haplotype network. I found a nonrandom association between haplotypes and life-history phenotypes. These results suggest that historical events responsible for population fragmentation may have also contributed to phenotypic shifts in life histories, both indirectly by limiting gene flow among populations and directly by altering the selective environments where populations persisted.     

Microsatellite markers for the June sucker (Chasmistes liorus mictus), Utah sucker (Catostomus ardens), and five other catostomid fishes of western North America

We developed and optimized five new microsatellite markers for the genetic management of the endangered June sucker. We report the cross‐amplification of these markers, and seven microsatellites previously developed for Klamath Basin suckers, in seven catostomid species of western North America. No linkage disequilibrium was detected between pairs of loci. Since most of these loci exhibited conserved priming sites, they may be useful for landscape‐scale studies of speciation and patterns of gene flow among multiple sucker lineages.     

Demography of an endangered,long-lived fish: Informing management options in the face of cyclic and stochastic climate variation

June sucker (Chasmistes liorus) is a long-lived, endangered fish endemic to Utah Lake, Utah. For several decades June sucker have failed to recruit sufficient numbers to the adult size classes such that the current wild population consists of a small number of old adults and it continues to decline. Vital rates of June sucker are influenced by climate-driven variation in lake level and inflow from the Provo River. We used population projection matrix modeling to assess effects of cyclic and stochastic environmental variation on population growth trajectories of June sucker in Utah Lake. The stable stage distribution is dominated by stage 1 individuals (93% of the total population) in contrast to the current situation where old age classes are the most abundant. Total population size is highly influenced by the stochastic component of climate variation; whereas, the adult population of June sucker closely tracks the systematic drought cycle. If changes in survival of larvae and juveniles can be coordinated such that positive changes in both parameters can occur somewhat simultaneously, then each parameter would only have to be increased by a factor of about 8.8 to achieve sustainable population growth (compared to a 77-fold increase for each parameter separately). Stochastic climatic variation has relatively little long-term effect on population growth. However, the multidecadal cyclic pattern of lake level and river discharge imposes a similar pattern on population growth rates of the June sucker, such that during some periods, populations decline even when the long-term trend is positive.     

Historic and recent age structure and growth of endangered Lost River and shortnose suckers in Upper Klamath Lake, Oregon

Seventy-four lapilli from Lost River suckers captured in Upper Klamath Lake in 1970 during a snag fishery on spawning adults and 192 lapilli from adults sacrificed from 2001–2006 were examined to determine age and growth parameters; lapilli from 165 shortnose suckers sacrificed from Upper Klamath Lake from 2001–2006 were also examined. Relative marginal distance analyses indicated that growth marks were annuli and formed in December–January. Lost River suckers from the historic collection were aged to 57 years, while Lost River and shortnose suckers from the recent collection were aged to 40 years and 24 years, respectively. Larger and older Lost River suckers were represented in the historic collection compared to the recent collection. Uncoupling of otolith length and fish length in Lost River suckers as well as a large spread in the predicted age- at-size for shortnose suckers precluded the ability to back-calculate size-at-age. Likelihood ratio tests indicated the growth model parameters were significantly different at both the sex and collection level. Growth in body length for both species appeared determinate in that growth was rapid until maturity, and then slowed over several years until growth in length was nearly nonexistent; a 650–700 mm Lost River sucker could be between 14 and 57 years old, while a 460 mm shortnose sucker could range from 12–24 years old. In contrast, while growth in body length slowed for both species, body mass continued to increase. This growth strategy, which is also found in other western lake suckers, may allow for more energy to be utilized for reproduction and help populations persist in spite of years of limited recruitment or recruitment failure.     

Differences in Species Composition and Feeding Ecology of Catostomid Fishes in Two Distinct Segments of the Missouri River, North Dakota, U.S.A.

In 1997 and 1998, we sampled the Missouri River, North Dakota to determine if anthropogenic disturbances had influenced catostomid species composition and feeding ecology. We compared two distinct river segments, the Missouri River between the mouth of the Yellowstone River and Lake Sakakawea (the Yellowstone–Sakakawea segment (YSS)), a moderately altered segment and the Missouri River between Garrison Dam and Lake Oahe (the Garrison–Oahe segment (GOS)), a highly altered segment. The segments exhibited greatly different sucker communities. Bigmouth buffalo, Ictiobus cyprinellus, smallmouth buffalo, Ictiobus bubalus, and river carpsucker, Carpiodes carpio, represented 94% of the sucker catch in the YSS, whereas in the GOS, white sucker, Catostomus commersoni, and longnose sucker, Catostomus catostomus, constituted 98% of the sucker catch. In the YSS, high zooplankton densities led to greater sucker zooplanktivory and food niche overlap than in the GOS. Intense anthropogenic disturbances to the GOS are associated with the differences in sucker species composition, prey density and composition, and sucker feeding ecology between the two segments.     

Comparison of electrofishing and trammel netting variability for sampling native fishes

The variability in size structure and relative abundance (CPUE; number of fish ≥200 mm total length, L_T, collected per hour of electrofishing or trammel netting) of three native Colorado River fishes, the endangered humpback chub Gila cypha, flannelmouth sucker Catostomus latipinnus and bluehead sucker Catostomus discobolus, collected from electrofishing and trammel nets was assessed to determine which gear was most appropriate to detect trends in relative abundance of adult fishes. Coefficient of variation (CV) of CPUE ranged from 210 to 566 for electrofishing and 128 to 575 for trammel netting, depending on season, diel period and species. Mean CV was lowest for trammel nets for humpback chub (P = 0·004) and tended to be lower for flannelmouth sucker (P = 0·12), regardless of season or diel period. Only one bluehead sucker >200 mm was collected with electrofishing. Electrofishing and trammel netting CPUE were not related for humpback chub (r = ?0·32, P = 0·43) or flannelmouth sucker (r = ?0·27, P = 0·46) in samples from the same date, location and hour set. Electrofishing collected a higher proportion of smaller (<200 mm L_T) humpback chub (P < 0·001), flannelmouth suckers (P < 0·001) and bluehead suckers (P < 0·001) than trammel netting, suggesting that conclusions derived from one gear may not be the same as from the other gear. This is probably because these gears fished different habitats, which are occupied by different fish life stages. To detect a 25% change in CPUE at a power of 0·9, at least 473 trammel net sets or 1918 electrofishing samples would be needed in this 8 km reach. This unattainable amount of samples for both trammel netting and electrofishing indicates that detecting annual changes in CPUE may not be practical and analysis of long‐term data or stock assessment models using mark‐recapture methods may be needed to assess trends in abundance of Colorado River native fishes, and probably other rare fishes as well.     

Ligulosis associated with mortality in largescale suckers

Investigations of a fish kill at an irrigation reservoir in the Deschutes River basin, Oregon, documented at least 153 dead largescale suckers Catostomus macrocheilus , all heavily infected with large Ligula intestinalis (Cestoda). Moribund fish nearshore were similarly infected, with no evidence of secondary disease or infection. Morbidity and mortality in all cases was probably due to severe mechanical damage to internal host organs resulting from infection with multiple, large L. intestinalis . The pathology of L. intestinalis documented in largescale sucker in this study contrasts sharply with previous accounts and may represent a new and emerging disease for largescale suckers.     

Finding a new form of the grayling Thymallus arcticus (thymallidae) in the basin of lake baikal

A dwarf form of the Arctic grayling Thymallus arcticus, inhabiting a group of small lakes at the origin of the Yakchii stream, which flows into the Verkhnyaya Angara River (basin of Lake Baikal), has been found in the northeastern watershed of the Baikal and Lena basins. The form is similar to upper Lena populations in body coloration and the pattern of dorsal fin. Our comparison by meristic characters indicated that the grayling of the Yakchii Lakes is more similar to the fish from Kutima River (basin of the upper courses of the Lena River) than to the black Baikal grayling T. arcticus baicalensis. The presence in Lake Baikal of a population similar to upper Lena graylings may have three causes: (1) possible drainage from Baikal to the pra-Lena via ancient valleys of the Barguzin and the Upper Angara; (2) appearance of transit zones as a result of glacial or tectonic events during the periods of watershed development between the Baikal and Lena basins; (3) the upper Lena grayling could be an endemic of the Baikal basin which was replaced by graylings penetrating from the Yenisei basin and remained in the form of relic populations in the upper courses of certain tributaries of the northern and northeastern parts of Lake Baikal.     

Effects of stand age and litter removal on the regeneration of Populus tremuloides

Abstract. The objective of this study was to demonstrate the regeneration by root suckering of Populus tremuloides (quaking aspen) along a successional sere at Lake Duparquet in NW Québec with six sites of different ages - 46, 74, 120, 143, 167 and 230 yr since last fire. In 1990, in each of the sites, one 20 m x 20 m plot was cut. In each plot, 4-m² plots were selected, both control and with litter removed. The number of suckers and their height were estimated for each plot in 1991, 1992 and 1993. Results showed that sucker production was not related to the number of P. tremuloides before the clearcut, but it was related to litter whereby increasing cover lowered the production. The number of suckers produced in the control plots did not show any significant difference along the successional sere for any of the three years. Scarified plots had more suckers than control plots, and this increase was more pronounced and significantly different for the younger stage. Regression analysis for the 1991 and 1992 data showed that sucker number in scarified plots decreased with increasing seral stage age. Mortality, after three growing seasons, was about 60% for all plots and was not affected by seral stage and treatment. We suggest that removal of litter in scarified plots exposed the roots to higher temperatures and caused wounds, both of which may have stimulated the production of suckers in young seral stages.     

Recent,small beginnings: genetic analysis suggests Catostomus rimiculus (Klamath smallscale sucker) in the Smith River,California, are introduced

Identification of introduced species can be important to understanding ecological systems and meeting conservation and management goals, but the process can be surprisingly challenging. The Klamath smallscale sucker Catostomus rimiculus seems likely to be native to the Smith River because the drainage separates two basins believed to be within the fish's native range, the Rogue and Klamath rivers. Further, C. rimiculus is broadly distributed in the Smith River, and the indigenous Dee-ni’ People of the Smith River have a unique word for sucker. Nonetheless, a historical survey of fishes that described C. rimiculus from the Rogue and Klamath rivers did not include C. rimiculus among the fishes of the Smith River. To determine whether the genetic structure of the Smith River C. rimiculus reflects expectations for a native sucker population, the authors of this study examined variation in microsatellite and mitochondrial genetic markers from the Smith River and surrounding drainages. The genetic analyses revealed a pattern consistent with extreme founder effects in Smith River C. rimiculus, as would be expected from a single introduction of six or fewer effective individuals. The sharing of a high-frequency haplotype between the Smith River and Klamath River that is not detected in the Rogue River suggests the Klamath River as the likely source for the introduction. The findings highlight that local-scale introductions can be easily overlooked because the newly established populations can appear to be parts of contiguous natural distributions.     

Postglacial dispersal of longnose suckers, Catostomus catostomus, in the Mackenzie and Yukon Drainages

The role played by stream transfers and reversals in the postglacial dispersal of fish populations in the Yukon and Mackenzie River drainages of Canada was analysed using genetic and meristic data from longnose sucker (Catostomus catostomus Forrester) populations. The hypothesis the suckers dispersed from the Yukon River into the Mackenzie River via the Peel River was evaluated. Meristic data (gill rakers and lateral line scales) were uninformative for this analysis. Genetic affinity between the Yukon River and upper Peel River populations, and between Mackenzie and lower Peel River populations is suggested by transferrin allele frequencies. These affinities support the view that longnose suckers inhabiting the middle and upper regions of the Peel River are derived from Beringian and Mississippian ancestors, while suckers in the lower Peel are descended from Mississippian stock. Evidence for dispersal by way of the Eagle River was not found. Affinities of sucker populations in other river systems in the region are also described.     

Genetic diversity and divergence among freshwater mussel (Anodonta) populations in the Bonneville Basin of Utah

Populations of the freshwater mussel genus Anodonta appear to be in a state of rapid decline in western North America, following a trend that unfortunately seems to be prevalent among these animals (Mollusca: Unionoida). Here we describe the patterns of molecular divergence and diversity among Anodonta populations in the Bonneville Basin, a large sub-basin of the Great Basin in western North America. Using amplified fragment length polymorphism (AFLP) analysis, we found a striking lack of nuclear diversity within some of these populations, along with a high degree of structuring among populations (FST = 0.61), suggesting post-Pleistocene isolation, due either to a long-term loss of hydrologic connectivity among populations or to more recent fish introductions. We also found evidence of recent hybridization in one of these populations, possibly mediated by fish-stocking practices. Using mitochondrial sequence data, we compared the Bonneville Basin populations to Anodonta in several other drainages in western North America. We found a general lack of resolution in these phylogenetic reconstructions, although there was a tendency for the Bonneville Basin Anodonta (tentatively A. californiensis) to cluster with A. oregonensis from the adjacent Lahontan Basin in Nevada. We recommend further investigation of anthropogenic factors that may be contributing to the decline of western Anodonta and a broad-scale analysis and synthesis of genetic and morphological variation among Anodonta in western North America.     

A molecular phylogeny of aquatic gastropods provides a new perspective on biogeographic history of the Snake River Region

Mitochondrial DNA sequences of aquatic gastropods of the subgenus Pyrgulopsis (Natricola) were analyzed to test a commonly accepted hypothesis concerning the early history of the Snake River in the northwestern US. Distributions of Natricola and other regional biota were previously used to infer that the Snake River flowed to the Pacific through southeastern Oregon and northern California during the Neogene prior to its capture by the Columbia River in the late Pliocene (2 Ma). A molecular phylogeny based on partial sequences of COI and NDI (1149 bp) indicates that the Natricola clade is restricted to the modern Snake-Columbia River Basin and the Oregon Lakes region whereas northern California populations previously assigned to this subgenus belong to other lineages. The Natricola clade is not deeply subdivided into Oregon Lakes and Snake River Basin units consistent with late Pliocene fragmentation of the hypothesized paleodrainage, but instead is shallowly structured and contains multiple transitions among these two geographic areas. The strongly supported sister relationship between Natricola and a species from northwest Nevada (P. imperialis) is consistent with a recent proposal that the ancestral Snake River did not flow through southeast Oregon but instead flowed south to the Humboldt River. Within the context of this hypothesis, the multiple transitions between the Snake River Basin and the Oregon Lakes region that occurred within Natricola may be attributed to a late Pleistocene connection between these areas that was unrelated to the early course of the Snake River.     

Morphological and molecular variation in the least madtom Noturus hildebrandi (Siluriformes: Ictaluridae), a Mississippi Embayment endemic: evidence for a cryptic lineage in the Hatchie River

Sheared principal component analysis of 40 morphometric characteristics measured for 146 individuals and relative frequencies of pigmentation patterns scored for 980 individuals of the least madtom Noturus hildebrandi, a diminutive catfish endemic to eastern lowland drainages of the Mississippi Embayment region of North America, suggested a clinal pattern of morphological variation extending across the range from north to south. DNA sequence data representing 90 individuals from the mitochondrial gene cytochrome b (cytb) analysed using Bayesian phylogenetic methods recovered four major haplotype clades, suggestive of a high degree of isolation by drainage. Individual gene trees of cytb and four additional nuclear loci as well as trees based on concatenated datasets of these genes consistently recovered a cryptic lineage of individuals from the Hatchie River drainage that is morphologically indistinguishable from surrounding populations. Gene‐tree analyses failed to recover a monophyletic N. hildebrandi with respect to Noturus baileyi. A coalescence‐based species tree analysis, however, did recover N. hildebrandi monophyly with high support, suggesting that relationships reflected in individual gene trees and concatenated datasets are in part artefacts of incomplete lineage sorting or an ancient introgressive event. Results are consistent with the hypothesis of an ancient connection between the Hatchie and Tennessee River systems. Current subspecific designations are of limited utility as they reflect morphological variation and are not entirely consistent with phylogeny. Discrepancies between the pattern of variation observed in the morphological and molecular data may be explained by recent local adaptation to individual stream conditions that masks deeper evolutionary divergences.     

Structural changes in gills of Lost River suckers exposed to elevated pH and ammonia concentrations

The Lost River sucker (Deltistes luxatus) is a federally listed, endangered fish that occurs primarily in Upper Klamath Lake-a hypereutrophic lake in southern Oregon, USA. A decline of the sucker population in the lake over the past few decades has been partly attributed to adverse water quality conditions, including elevated pH and ammonia concentrations that occur during summer cyanobacterial blooms. We quantitatively analyzed structural changes in gills of larval Lost River suckers after they were exposed to elevated pH and ammonia concentrations for 30 d. Exposure to pH as high as 10 caused no observed structural changes. However, lamellar thickness and O(2) diffusion distance increased significantly (P<0.05) at ammonia concentrations that did not significantly decrease survival, growth, whole-body ion concentrations, or swimming performance. Additionally, we qualitatively observed increases in the frequency of hyperplasic and hypertrophic mucous cells, tissue damage, epithelial lifting, and infiltration of white blood cells into paracellular lymphatic spaces at the highest sublethal ammonia concentration. These observed gill changes typically indicate compromised respiratory and ionoregulatory capacity, although such effects were not manifested in the assays we performed. Regardless, these structural gill changes appear to be a more sensitive indicator of exposure to elevated ammonia concentrations than are more traditional sublethal indices. Therefore, gill histopathology might be a relevant early-warning monitoring tool of the health of Lost River suckers in Upper Klamath Lake, and other species in similar eutrophic systems.     

Comparison between clonal and sexual plantlets of Detarium microcarpum Guill. & Perr., a savanna tree species in Burkina Faso

An understanding of the proportion of true seedlings, seedling sprouts and root suckers in the forest is essential for directing the genetic composition of the future crop. We conducted a study to determine the difference between these plantlets of Detarium microcarpum based on morphological characters and carbohydrate contents in leaves and roots. For individuals ≤50 cm in height, root suckers had the highest values for height, stem length, internode number, root diameter, rachis length and leaflet number. The concentrations of starch and total nonstructural carbohydrates in the roots of seedling sprouts were superior. Plantlets did not differ in the concentration of leaf carbohydrates. For individuals >50 cm in height, root suckers had larger values for stem length, root diameter, leaflet length and width. Roots of seedling sprouts showed higher concentrations of soluble sugars and total soluble sugars. True seedlings were distinguished from seedling sprouts and root suckers using all morphological traits except collar diameter and leaflet number. Root suckers and seedling sprouts showed a closer morphological resemblance; thus resulted in slightly more than 50% discrimination success. In conclusion, discrimination between seedling sprouts and root suckers appeared to be more difficult than between true seedlings and clonal plantlets.     

Morphological variation and taxonomic status of Altai osmans, Oreoleuciscus, from the upper reaches of the Ob River system

The morphology of the endemic Altai osmans Oreoleuciscus spp. (Cyprinidae), from headwaters of the Ob tributaries in the Altai Republic (Russia), Terekhol Lake in the Tuva Republic (Russia), and the Chovd Gol River drainage and related bodies of water (Mongolia), where they are one of only four fish species occurring in most bodies of water were examined. Their high degree of morphological variation makes species determination problematical. Analysis of six meristic counts and principal component analysis (PCA) of 21 morphological measurements discriminated two species, Oreoleuciscus potanini and O. humilis . The results suggest that only O. potanini occurs in the upper reaches of the Ob system. The systematics and biogeography of Oreoleuciscus are discussed.     

Neglected taxonomy of rare desert fishes: congruent evidence for two species of leatherside chub

Conservation biologists rely heavily on taxonomy to set the scope for biological monitoring and recovery planning of rare or threatened species. Yet, taxonomic boundaries are seldom evaluated as falsifiable hypotheses that can be statistically tested. Here, we examine species boundaries in leatherside chub (Teleostei, Cyprinidae), an imperiled desert fish native to the Bonneville Basin and upper Snake River drainages of western North America. Recent molecular data hint that this fish could be composed of two distinct taxa that are geographically separated into northern and southern species. To formally test this hypothesis, we evaluated leatherside chub using several different categories of species concepts, including criteria dependent on phylogenetic, morphological, and ecological data. We found that leatherside chub is composed of two reciprocally monophyletic clades (candidate species) characterized by numerous fixed genetic differences for both mitochondrial and nuclear DNA markers; mtDNA sequence divergence between the two clades approached 8%. The candidate species also showed significant differences in cranial shape, revealed by morphometric analysis. Finally, controlled growth and foraging experiments using representative populations from each clade show that candidate species appear to be locally adapted to the thermal environments where they now occur. Combined, these three lines of evidence support the hypothesis that leatherside chub is composed of two species. Moreover, all lines of evidence place these two species within the genusLepidomeda, a group consisting of three additional species of endangered spinedace fishes, and one extinct species, all native to the Colorado River system. Hence, we elevate the two clades of leatherside chub to distinct species status (Lepidomeda copeiin the north andL. aliciaein the south), and argue that each warrants independent conservation and recovery action.