嵩草属植物的系统学研究

嵩草属隶属于莎草科苔草亚科苔草族，主要分布于北半球温带地区，少数种类为环北极分布，有一种产于泰国清迈，一种产于苏门达腊岛北部高山。本‘文在形态学、微形态学、解剖学和胚胎学研究的基础上，对嵩草属植物进行了全面的分类学修订，根据属内各类群之间的系统演化关系建立了一个新的属下分类系统，确认了世界范围内53种3亚种嵩草属植物，并做出5个种上等级的新组合，描述了一个新亚组。 作者研究了国内外14家标本馆（室）的3000多份腊叶标本并进行多次野外的实地观察，对嵩草属植物的形态学性状进行了详细的比较和分析，评价了它们的系统学价值及其演化趋势。在嵩草属中，花序是由小穗排列成的圆锥花序或穗状花序，花序各部分的形态性状是种及种上等级分类的基础;花序的演化趋势是由圆锥花序到穗状花序，小穗是从由数朵雄花与1朵雌花组成简化到由1朵雌花组成。但是，花序和小穗由复杂到简单的进化在嵩草属中平行地发生于不同的类群中。先出叶的性状状态是分种的主要特征之一，通常认为边缘开裂的先出叶是原始的，边缘合生而为囊状的先出叶是进化的。同样，先出叶由开裂到合生的进化也是多次发生的。此外，根状茎、秆、叶鞘、叶片、柱头及小坚果等的性状状态对于种及种上等级的分类都具有重要的意义。 应用扫描电子显微镜对38种（或亚种）嵩草属植物的小坚果表面进行了观察，证明小坚果纹饰在种及种上等级的分类中具有重要的参考价值，并能揭示种及种下等级的亲缘关系。例如，分布于喜马拉雅东部至横断山地区的3种植物，K．clarkeana、K．curvata和K．fragilis外部形态非常相似，难于区分，而其小坚果的微形态特征却可以提供3种之间关系的证据。K．clarkeana与K．fragilis果实表面的特征完全一致，且与其它植物有显著区别，应为同种植物;K．curvata与它们明显不同，也与其它种有较大差异，应为独立的种。K．gramini folia，K．cercostach ys和K. nepalensis果实表面纹饰具有一些共同的特征，说明它们之间的亲缘关系较近。K．filicina和K．duthiei也存在同样的情形。 通过对秆和叶片的横切面和表皮的解剖学观察比较，发现嵩草属植物秆的横切面表现出由三角形到圆形的一系列变化。秆的横切面明显地分两个区域，中部的髓由较大而无色的细胞组成，其中心常碎裂形成大的气腔;外围的绿色部分，由绿色组织及分布其中的气腔和外韧维管束及与其相伴的厚壁组织组成。秆的表皮与叶片下表皮非常相象。叶片横切面的外形为V形、新月形或半圆形。V形的叶片具有明显发育的中脉并且在远轴面凸起，形成脊;新月形和半圆形的叶片中脉发育不明显，也无脊。叶片的表皮细胞均为长方形，垂周壁波纹状;平列型的气孔器纵向成行排列，多局限于下表皮;上表皮近边缘及脉附近的细胞常常在细胞的一端形成乳突。秆和叶片横切面的形态对于分种及种上等级的划分具有参考价值。 胚胎学研究表明小孢子、胚囊和胚的发育与莎草科其它类群一致。花粉为假单体花粉( pseudomonad)，成熟花粉三核。胚珠倒生，厚珠心，双层珠被，珠孑L由内珠被形成。胚囊的发育为蓼型，原胚的发育为柳叶菜型灯芯草变型。首次观察到，在大孢子四分体时期，合点端和珠孔端两个大孢子细胞开始时体积都增大，而中部两个很快退化，稍后珠孔端一个也退化，合点端一个为功能大孢子，发育成为胚囊。根据胚胎学证据，不支持将嵩草属与苔草族一起另立为嵩草科。 嵩草属中较原始的一个亚属subg. Compositae主要分布于西喜马拉雅至横断山地区，还有一种见于泰国，一种产于苏门达腊，而后2种植物还具有一些最原始的形态性状。结合地史的变化推测，嵩草属可能在第三纪早期起源于古地中海的东部和北部。 根据形态学和解剖学性状的分析表明，许多性状在嵩草属中是平行演化的，如花序和小穗由复杂到简单、秆由圆柱形到三棱形、叶片横切面由V形到半圆形等。该属的属下分类应该追溯这些平行的演化线，而不能像以前的分类那样，将它们横向地划分为几个组或亚属。作者认为嵩草属有3个大的进化分支，据此将其划分为3个亚属。Subg. Compositae，12种，是较原始的一个分支。叶近基生，叶片扁平;花序多疏松圆锥状，少穗状，苞片多为叶状;小穗两性到单性，先出叶多为囊状，少边缘分离，退化小穗轴明显、扁平、较长。Subg. Blysmocarex，仅2种，是较早分化而相对隔离的一个分支。根状茎匍匐状;花序由圆锥状到穗状，小穗两性或单性;柱头2。Subg. Kobresia，种类最多。叶片扁平或内卷;秆三棱形到圆柱形;花序紧密，复杂到简单，苞片不为叶状;小穗两性或单性，先出叶由开裂到合生，退化小穗轴较小而不显著。根据该亚属呈现出的不同的性状演化系列，可以分为3个组。Sect. Kobresia花序圆锥状至穗状，小穗多为两性，少为单性，先出叶边缘分离。含3个亚组：subsect. Kobresia，8种2亚种．植株纤细，秆与叶均为丝状;subsect. Royleanae，8种l亚种，植株较粗壮，叶片扁平或对折;subsect. Sibiri-cae，4种，秆较粗，叶片内卷。Sect. Psmmostachys，仅2种，小穗两性，先出叶完全合生为囊状。Sect．Hemicarex花序一般为穗状，稀圆锥状，小穗多为单性，少两性。分为四个亚组：subsect. Forexeta，6种，叶片内卷或对折，先出叶线形，边缘分离或合生;subsect. Chlorostachys，3种，叶片扁平，小穗两性;subsect. Holmia，4种，叶片扁平，小穗单性;subsect．Utriculatae，5种，叶片丝状，先出叶完全合生为囊状，不为线形。嵩草属的属下分类纲要如下： Subgenus 1. Compositae (Clarke) Kukkonen Type: K. laxa Nees. Subgenus 2. Blysmocarex (Ivanova) S. R. Zhang Type: K. macrantha Boeck. Subgenus 3. Kobresia Section 1. Kobresia Subsection 1. Kobresia Type: K. simpliciuscula (Wahl. ) Mack. Subsection 2. Royleanae (Ivanova) S. R. Zhang Type: K. royleana (Nees) Boeck. Subsection 3. Sibiricae (Ivanova) Egorova Type: K. sibirica (Turcz. ex Ledeb. ) Boeck. Section 2. Psmmostachys Ivanova Type: K. robusta Maxim. Section 3. Hemicarex (Bentham) Clarke Subsection 4. Forexeta (Raffin. ) S. R. Zhang Type: K. cercostachys (Franch. ) C. B. Clarke. Subsection 5. Chlorostachys (Ivanova) S. R. Zhang Type: K. duthiei C B. Clarke. Subsection 6. Holmia (Boern. ) S. R. Zhang Type: K. esenbeckii (Kunth) Noltie. Subsection 7. Utriculatae S. R. Zhang Type: K. prainii Kukenthal.

Year-to-year changes of vertical temperature distribution in the California Current region: 1954 to 1986

Studies by Enfield and Allen (1980), McLain et al (1985), and others have shown that anomalously warm years in the northern coastal California Current correspond to El Niño conditions in the equatorial Pacific Ocean. Ocean model studies suggest a mechanical link between the northern coastal California Current and the equatorial ocean through long waves that propagate cyclonically along the ocean boundary (McCreary 1976; Clarke 1983; Shriver et al 1991). However, distinct observational evidence of such an oceanic connection is not extensive. Much of the supposed El Niño variation in temperature and sea level data from the coastal California Current region can be associated with the effects of anomalously intense north Pacific atmospheric cyclogenesis, which is frequently augmented during El Niño years (Wallace and Gutzler 1981; Simpson 1983; Emery and Hamilton 1984). This study uses time series of ocean temperature data to distinguish between locally forced effects, initiated by north Pacific atmospheric changes, and remotely forced effects, initiated by equatorial Pacific atmospheric changes related to El Niño events.

A biogeographic assessment of the Channel Islands National Marine Sanctuary: a review of boundary expansion concepts for NOAA’s National Marine Sanctuary Program

The priority management goal of the National Marine Sanctuaries Program (NMSP) is to protect marine ecosystems and biodiversity. This goal requires an understanding of broad-scale ecological relationships and linkages between marine resources and physical oceanography to support an ecosystem management approach. The Channel Islands National Marine Sanctuary (CINMS) is currently reviewing its management plan and investigating boundary expansion. A management plan study area (henceforth, Study Area) was described that extends from the current boundary north to the mainland, and extends north to Point Sal and south to Point Dume. Six additional boundary concepts were developed that vary in area and include the majority of the Study Area. The NMSP and CINMS partnered with NOAA’s National Centers for Coastal Ocean Science Biogeography Team to conduct a biogeographic assessment to characterize marine resources and oceanographic patterns within and adjacent to the sanctuary. This assessment includes a suite of quantitative spatial and statistical analyses that characterize biological and oceanographic patterns in the marine region from Point Sal to the U.S.-Mexico border. These data were analyzed using an index which evaluates an ecological “cost-benefit” within the proposed boundary concepts and the Study Area. The sanctuary resides in a dynamic setting where two oceanographic regimes meet. Cold northern waters mix with warm southern waters around the Channel Islands creating an area of transition that strongly influences the regions oceanography. In turn, these processes drive the biological distributions within the region. This assessment analyzes bathymetry, benthic substrate, bathymetric life-zones, sea surface temperature, primary production, currents, submerged aquatic vegetation, and kelp in the context of broad-scale patterns and relative to the proposed boundary concepts and the Study Area. Boundary cost-benefit results for these parameters were variable due to their dynamic nature; however, when analyzed in composite the Study Area and Boundary Concept 2 were considered the most favorable. Biological data were collected from numerous resource agencies and university scientists for this assessment. Fish and invertebrate trawl data were used to characterize community structure. Habitat suitability models were developed for 15 species of macroinvertebrates and 11 species of fish that have significant ecological, commercial, or recreational importance in the region and general patterns of ichthyoplankton distribution are described. Six surveys of ship and plane at-sea surveys were used to model marine bird diversity from Point Arena to the U.S.-Mexico border. Additional surveys were utilized to estimate density and colony counts for nine bird species. Critical habitat for western snowy plover and the location of California least tern breeding pairs were also analyzed. At-sea surveys were also used to describe the distribution of 14 species of cetaceans and five species of pinnipeds. Boundary concept cost-benefit indices revealed that Boundary Concept 2 and the Study Area were most favorable for the majority of the species-specific analyses. Boundary Concept 3 was most favorable for bird diversity across the region. Inadequate spatial resolution for fish and invertebrate community data and incompatible sampling effort information for bird and mammal data precluded boundary cost-benefit analysis.

The Southeast Bering Sea Ecosystem: implications for marine resource management (Final Report: Southeast Bering Sea Carrying Capacity).

Southeast Bering Sea Carrying Capacity (SEBSCC, 1996–2002) was a NOAA Coastal Ocean Program project that investigated the marine ecosystem of the southeastern Bering Sea. SEBSCC was co-managed by the University of Alaska Fairbanks, NOAA Alaska Fisheries Science Center, and NOAA Pacific Marine Environmental Laboratory. Project goals were to understand the changing physical environment and its relationship to the biota of the region, to relate that understanding to natural variations in year-class strength of walleye pollock (Theragra chalcogramma), and to improve the flow of ecosystem information to fishery managers. In addition to SEBSCC, the Inner Front study (1997–2000), supported by the National Science Foundation (Prolonged Production and Trophic Transfer to Predators: Processes at the Inner Front of the S.E. Bering Sea), was active in the southeastern Bering Sea from 1997 to 1999. The SEBSCC and Inner Front studies were complementary. SEBSCC focused on the middle and outer shelf. Inner Front worked the middle and inner shelf. Collaboration between investigators in the two programs was strong, and the joint results yielded a substantially increased understanding of the regional ecosystem. SEBSCC focused on four central scientific issues: (1) How does climate variability influence the marine ecosystem of the Bering Sea? (2) What determines the timing, amount, and fate of primary and secondary production? (3) How do oceanographic conditions on the shelf influence distributions of fish and other species? (4) What limits the growth of fish populations on the eastern Bering Sea shelf? Underlying these broad questions was a narrower focus on walleye pollock, particularly a desire to understand ecological factors that affect year-class strength and the ability to predict the potential of a year class at the earliest possible time. The Inner Front program focused on the role of the structural front between the well-mixed waters of the coastal domain and the two-layer system of the middle domain. Of special interest was the potential for prolonged post-spring-bloom production at the front and its role in supporting upper trophic level organisms such as juvenile pollock and seabirds. Of concern to both programs was the role of interannual and longer-term variability in marine climates and their effects on the function of sub-arctic marine ecosystems and their ability to support upper trophic level organisms.

Skin lesions on common bottlenose dolphins (Tursiops truncatus) from three sites in the Northwest Atlantic, USA

Skin disease occurs frequently in many cetacean species across the globe; methods to categorize lesions have relied on photo-identification (photo-id), stranding, and bycatch data. The current study used photo-id data from four sampling months during 2009 to estimate skin lesion prevalence and type occurring on bottlenose dolphins (Tursiops truncatus) from three sites along the southeast United States coast [Sarasota Bay, FL (SSB); near Brunswick and Sapelo Island, GA (BSG); and near Charleston, SC (CHS)]. The prevalence of lesions was highest among BSG dolphins (P=0.587) and lowest in SSB (P=0.380), and the overall prevalence was significantly different among all sites (p<0.0167). Logistic regression modeling revealed a significant reduction in the odds of lesion occurrence for increasing water temperatures (OR=0.92; 95%CI:0.906-0.938) and a significantly increased odds of lesion occurrence for BSG dolphins (OR=1.39; 95%CI:1.203-1.614). Approximately one-third of the lesioned dolphins from each site presented with multiple types, and population differences in lesion type occurrence were observed (p<0.05). Lesions on stranded dolphins were sampled to determine the etiology of different lesion types, which included three visually distinct samples positive for herpesvirus. Although generally considered non-fatal, skin disease may be indicative of animal health or exposure to anthropogenic or environmental threats, and photo-id data provide an efficient and cost-effective approach to document the occurrence of skin lesions in free-ranging populations.

Anti-grazing properties of the toxic dinoflagellate Karlodinium veneficum during predator-prey interactions with the copepod Acartia tonsa

Karlodinium veneficum (syn. Karlodinium micrum, Bergholtz et al. 2006; J Phycol 42:170–193) is a small athecate dinoflagellate commonly present in low levels in temperate, coastal waters. Occasionally, K. veneficum forms ichthyotoxic blooms due to the presence of cytotoxic, hemolytic compounds, putatively named karlotoxins. To evaluate the anti-grazing properties of these karlotoxins, we conducted food removal experiments using the cosmopolitan copepod grazer Acartia tonsa. Wild-caught, adult female A. tonsa were exposed to 6 monoalgal or mixed algal diets made using bloom concentrations of toxic (CCMP 2064) and non-toxic (CSIC1) strains of K. veneficum. Ingestion and clearance rates were calculated using the equations of Frost (1972). Exposure to the toxic strain of K. veneficum did not contribute to an increased mortality of the copepods and no significant differences in copepod mortality were found among the experimental diets. However, A. tonsa had significantly greater clearance and ingestion rates when exposed to a monoalgal diet of the non-toxic strain CSIC1 than when exposed to the monoalgal diet of toxic strain CCMP 2064 and mixed diets dominated by this toxic strain. These results support the hypothesis that karlotoxins in certain strains of K. veneficum deter grazing by potential predators and contribute to the formation and continuation of blooms.

Radiometric validation of age, growth, and longevity for the blackgill rockfish (Sebastes melanostomus)

As nearshore fish populations decline, many commercial fishermen have shifted fishing effort to deeper continental slope habitats to target fishes for which biological information is limited. One such fishery that developed in the northeastern Pacific Ocean in the early 1980s was for the blackgill rockfish (Sebastes melanostomus), a deep-dwelling (300−800 m) species that congregates over rocky pinnacles, mainly from southern California to southern Oregon. Growth zone-derived age estimates from otolith thin sections were compared to ages obtained from the radioactive disequilibria of 210Pb, in relation to its parent, 226Ra, in otolith cores of blackgill rockfish. Age estimates were validated up to 41 years, and a strong pattern of agreement supported a longevity exceeding 90 years. Age and length data fitted to the von Bertalanffy growth function indicated that blackgill rockfish are slow-growing (k= 0.040 females, 0.068 males) and that females grow slower than males, but reach a greater length. Age at 50% maturity, derived from previously published length-at-maturity estimates, was 17 years for males and 21 years for females. The results of this study agree with general life history traits already recognized for many Sebastes species, such as long life, slow growth, and late age at maturation. These traits may undermine the sustainability of blackgill rockfish populations when heavy fishing pressure, such as that which occurred in the 1980s, is applied.

A model for assessing the likelihood of self-sustaining populations resulting from commercial production of triploid Suminoe oysters (Crassostrea ariakensis) in Chesapeake Bay

Culture of a non-native species, such as the Suminoe oyster (Crassostrea ariakensis), could offset the harvest of the declining native eastern oyster (Crassostrea virginica) fishery in Chesapeake Bay. Because of possible ecological impacts from introducing a fertile non-native species, introduction of sterile triploid oysters has been proposed. However, recent data show that a small percentage of triploid individuals progressively revert toward diploidy, introducing the possibility that Suminoe oysters might establish self-sustaining populations. To assess the risk of Suminoe oyster populations becoming established in Chesapeake Bay, a demographic population model was developed. Parameters modeled were salinity, stocking density, reversion rate, reproductive potential, natural and harvest-induced mortality, growth rates, and effects of various management strategies, including harvest strategies. The probability of a Suminoe oyster population becoming self-sustaining decreased in the model when oysters are grown at low salinity sites, certainty of harvest is high, mini-mum shell length-at-harvest is small, and stocking density is low. From the results of the model, we suggest adopting the proposed management strategies shown by the model to decrease the probability of a Suminoe oyster population becoming self-sustaining. Policy makers and fishery managers can use the model to predict potential outcomes of policy decisions, supporting the ability to make science-based policy decisions about the proposed introduction of triploid Suminoe oysters into the Chesapeake Bay.