The Bornean Lithocarpus B1. section Synaedrys (Lindl.) Barnett (Fagaceae): its circumscription and description of a new species

The Bomean species of Lithocarpus B1. section Synaedrys edl.) Barnett (Fagaceae) are discussed in terms of their unifying characteristics, and keys to the species are provided. A novel morphological feature present in the genus, where the seed in the mature fruit is enclosed by a lignified and thickened receptacle, is illustrated. A new species, L palungensis Cannon & Manos, is described. It can be distinguished from L. pulcher (King) Markgr. by its submontane distribution, prominent reflexed cupular spines, present at all stages, and broadly oblong leaves with scattered open fasciculate trichomes on the abaxial surface. Foliar trichome types present within the section are also described. Distinctiveness of fruit wall and receptacle shape, as described by two-dimensional elliptic Fourier transformation, is demonstrated by principal components analysis.     

Towards a molecular phylogeny ofApiaceae subfamilyApioideae: Additional information from nuclear ribosomal DNA ITS sequences

Evolutionary relationships among 116 representatives (80 genera) ofApiaceae (Umbelliferae) subfam.Apioideae were investigated by comparative sequencing of the two internal transcribed spacers of the 18S–26S nuclear ribosomal DNA repeat. The resultant phylogenies, inferred using maximum parsimony and neighbor-joining methods, clarified the relationships of several genera whose phylogenetic placements have heretofore been problematic. Comparisons between the phylogenies inferred and the distribution of several phytochemical (coumarins, flavonoids, and phenylpropenes) and morphological (stomates, pollen, and cotyledonary shape) characters were also made, revealing that many of these characters (like those morphological and anatomical characters of the fruit) are highly homoplastic. It is not surprising then that systems of classification ofApioideae based on these characters, particularly with regard to tribal and subtribal designations and relationships, are unsatisfactory. The results of recent serological investigations of the subfamily support several relationships proposed herein using molecular data.     

Evidence for a trade-off strategy in stone oak (Lithocarpus) seeds between physical and chemical defense highlights fiber as an important antifeedant

Trees in the beech or oak family (Fagaceae) have a mutualistic relationship with scatter-hoarding rodents. Rodents obtain nutrients and energy by consuming seeds, while providing seed dispersal for the tree by allowing some cached seeds to germinate. Seed predation and caching behavior of rodents is primarily affected by seed size, mechanical protection, macronutrient content, and chemical antifeedants. To enhance seed dispersal, trees must optimize trade-offs in investment between macronutrients and antifeedants. Here, we examine this important chemical balance in the seeds of tropical stone oak species with two substantially different fruit morphologies. These two distinct fruit morphologies in Lithocarpus differ in the degree of mechanical protection of the seed. For 'acorn' fruit, a thin exocarp forms a shell around the seed while for 'enclosed receptacle' (ER) fruit, the seed is embedded in a woody receptacle. We compared the chemical composition of numerous macronutrient and antifeedant in seeds from several Lithocarpus species, focusing on two pairs of sympatric species with different fruit morphologies. We found that macronutrients, particularly total non-structural carbohydrate, was more concentrated in seeds of ER fruits while antifeedants, primarily fibers, were more concentrated in seeds of acorn fruits. The trade-off in these two major chemical components was more evident between the two sympatric lowland species than between two highland species. Surprisingly, no significant difference in overall tannin concentrations in the seeds was observed between the two fruit morphologies. Instead, the major trade-off between macronutrients and antifeedants involved indigestible fibers. Future studies of this complex mutualism should carefully consider the role of indigestible fibers in the foraging behavior of scatter-hoarding rodents.     

Habitat transitions alter the adaptive landscape and shape phenotypic evolution in needlefishes (Belonidae)

Habitat occupancy can have a profound influence on macroevolutionary dynamics, and a switch in major habitat type may alter the evolutionary trajectory of a lineage. In this study, we investigate how evolutionary transitions between marine and freshwater habitats affect macroevolutionary adaptive landscapes, using needlefishes (Belonidae) as a model system. We examined the evolution of body shape and size in marine and freshwater needlefishes and tested for phenotypic change in response to transitions between habitats. Using micro‐computed tomographic (µCT) scanning and geometric morphometrics, we quantified body shape, size, and vertebral counts of 31 belonid species. We then examined the pattern and tempo of body shape and size evolution using phylogenetic comparative methods. Our results show that transitions from marine to freshwater habitats have altered the adaptive landscape for needlefishes and expanded morphospace relative to marine taxa. We provide further evidence that freshwater taxa attain reduced sizes either through dwarfism (as inferred from axial skeletal reduction) or through developmental truncation (as inferred from axial skeletal loss). We propose that transitions to freshwater habitats produce morphological novelty in response to novel prey resources and changes in locomotor demands. We find that repeated invasions of different habitats have prompted predictable changes in morphology.     

Phylogeny of Steinernema travassos, 1927 (Cephalobina: Steinernematidae) inferred from ribosomal DNA sequences and morphological characters

Entomopathogenic nematodes in Steinernema, together with their symbiont bacteria Xenorhabdus, are obligate and lethal parasites of insects that can provide effective biological control of some important lepidopteran, dipteran, and coleopteran pests of commercial crops. Phylogenetic relationships among 21 Steinernema species were estimated using 28S ribosomal DNA (rDNA) sequences and morphological characters. Sequences of the rDNA internal transcribed spacers were obtained to provide additional molecular characters to resolve relationships among Steinernema carpocapsae, Steinernema scapterisci, Steinernema siamkavai, and Steinernema monticolum. Four equally parsimonious trees resulted from combined analysis of 28S sequences and 22 morphological characters. Clades inferred from analyses of molecular sequences and combined datasets were primarily reliably supported as assessed by bootstrap resampling, whereas those inferred from morphological data alone were not. Although partially consistent with some traditional expectations and previous phylogenetic studies, the hypotheses inferred from molecular evidence, and those from combined analysis of morphological and molecular data, provide a new and comprehensive framework for evaluating character evolution of steinernematids. Interpretation of morphological character evolution on 6 trees inferred from sequence data and combined evidence suggests that many structural features of these nematodes are highly homoplastic, and that some structures previously used to hypothesize relationships represent ancestral character states.     

中国飘拂草属植物果皮微形态

利用扫描电子显微镜对国产莎草科飘拂草属4组18系42种1变种植物果皮微形态特征进行观察,并作了系统描述。研究表明,飘拂草属植物在纹饰类型,超微结构等方面存在着明显的种间差异,具有分类学意义。根据纹饰及微形态特征的不同,支持将褐鳞飘拂草和知风飘拂草各自作为独立的种。依照果皮纹饰的差异,飘拂草属可分为4种类型(1)瘤(疣)-网状复合纹饰;(2)瘤(疣)状纹饰;(3)网状纹饰;(4)脊-疣状复合纹饰。其中(1)和(3)类型的纹饰根据网脊曲直不同各自又可分为2个亚型。     

Molecular data reveal convergence in fruit characters used in the classification of Thlaspi s. I. (Brassicaceae)

Phylogenetic relationships of 18 Thlaspi s.l. species were inferred from nuclear ribosomal internal transcribed spacer (ITS) sequence data. These species represent all sections of the basic classification system of Schulz primarily based on fruit characters. The molecular phylogeny supported six clades that are largely congruent with species groups recognized by Meyer on the basis of differences in seed coat anatomy, i.e. Thlaspi s. s., Thlaspkeras, Moccaea {Raparia included), Microthhspi, Vania and Neurotropy. Some of these lineages include species which are morphologically diverse in fruit shape (e.g. Thlaspi s. s.: T. arvense - fruits broadly winged, T. ceratocarpum - fruits with prominent horns at apex, T. alliaceum - fruits very narrowly winged). Furthermore, the same fruit shape type is distributed among different clades. For instance, fruits with prominent horns at apex are found in Thlaspi s. s. ( T. ceratocarpum) and Thlaspiceras (T oxyceras). These results clearly indicate convergence in fruit characters previously used for sectional classification in Thlaspi s. l.     

Grazers, shredders and filtering carnivores--the evolution of feeding ecology in Drusinae (Trichoptera: Limnephilidae): insights from a molecular phylogeny

We examined the phylogenetic relationships between species and genera within the caddisfly subfamily Drusinae (Trichoptera: Limnephilidae) using sequence data from two mitochondrial loci (cytochrome oxidase 1, large subunit rRNA) and one nuclear gene (wingless). Sequence data were analysed for 28 species from five genera from the subfamily. We analysed individual and combined data sets using a Bayesian Markov Chain Monte Carlo and a maximum parsimony approach and compared the performance of each partition for resolving phylogenetic relationships at this level. In terms of resolution and phylogenetic utility wingless outperformed the two mitochondrial gene partitions. Using both Shimodaira-Hasegawa and expected likelihood weights tests we tested several hypotheses of relationships previously inferred based on adult morphological characters. The data did not support the generic concept, or many previously proposed species groupings, based on adult morphology. In contrast, the molecular data correlated with the morphology and feeding ecology of larvae. Using Bayesian ancestral character state reconstructions we inferred the evolution of feeding ecology and relevant larval morphological characters. Our analyses showed that within the subfamily Drusinae two derived feeding types evolved. One of these--grazing epilithic algae--is otherwise unusual in the Limnephilidae and may have promoted the high degree of diversity in the Drusinae.     

4种形态类型诸葛菜的等位酶变异及遗传多样性分析

以采自江苏省南京市4个样点的4种不同形态类型(紫花毛果、紫花光果、白花光果和白花毛果)诸葛菜〔Orychophragmus violaceus(L.)O.E.Schulz〕63个单株为研究对象,采用不连续聚丙烯酰胺凝胶电泳技术(SDS-PAGE)对过氧化物酶同工酶(POD)、乙醇脱氢酶(ADH)、谷氨酸脱氢酶(GDH)、超氧化物歧化酶(SOD)和多酚氧化酶(PPO)酶谱进行了分析,并基于5个等位酶的酶谱分析结果对4种形态类型诸葛菜的遗传多样性进行了研究。结果表明:5个等位酶系统共包含12个位点32个等位基因,其中POD-1、POD-4、POD-5、POD-6、ADH-1、PPO-1、SOD-1和SOD-2为多态性位点;除具有共有谱带外,不同类型诸葛菜还具有各自特有的酶谱特征,其中,GDH-2的b带和c带分别是紫花类型和白花类型的特有谱带。紫花毛果、紫花光果和白花毛果类型的多态位点百分率(PPL)均为58.33%,而白花光果的PPL为66.67%;紫花毛果、紫花光果、白花光果和白花毛果每个位点的平均等位基因数分别为2.42、2.25、2.42和1.83,平均期望杂合度分别为0.35、0.30、0.38和0.29。4种形态类型诸葛菜的总基因多样度平均值为0.61,类型内基因多样度平均值为0.49,明显大于类型间基因多样度平均值(0.12);类型间的基因分化系数平均值为0.195,表明总基因多样性的80.5%来源于类型内。紫花光果和白花毛果类型间的遗传一致度最低(0.724 4)且遗传距离最大(0.322 5),而2个白花类型间的遗传一致度最高(0.954 1)且遗传距离最小(0.047 1)。研究结果显示:诸葛菜的种内变异程度很大、遗传分化程度较高,但在各类型内具有较高的遗传相似性。     

Population structure and local differentiation in the delicate loach, Niwaella delicata, as revealed by mitochondrial DNA and morphological analyses

Population structures of the delicate loach, Niwaella delicata, were inferred from morphology and restriction fragment length polymorphism (RFLP) analysis of part of the mitochondrial DNA (mtDNA) of 25 populations, representing the species range in central Honshu Island. The existence of two types of morphological variation corresponding to regional distributions, the "Pacific slope type" and "Sea of Japan slope type," has been known in N. delicata. Our morphological reexamination of the two types revealed some discrepancies in their distribution pattern. Therefore, we reclassified two new color types corresponded to their distribution areas as "gathered spots type (G type)" and "scattered spots type (S type)," respectively. The present classification of G and S types is closely related to the mtDNA divergence pattern. The current analysis also indicated that each G and S type population was further divided into two genetic groups, corresponding to geographic proximity. In spite of marked morphological differentiation, the genetic diversity between G and S type populations (1.153%) was comparable only to that reported for intraspecific levels in most freshwater fishes. Moreover, in the population of which the color patterns of all fish were characterized to the S type, mtDNA haplotypes corresponding to G and S types were sympatrically detected. This result indicates secondary contact between the two type populations and the possibility that they are not reproductively isolated. Received: June 11, 1999 / Revised: September 30, 2000 / Accepted: January 16, 2001     

果皮和种皮微形态特征在杜鹃花属系统学研究中的应用

采用扫描电子显微镜对国产杜鹃花属Rhododendron6个亚属的代表种和近缘类群杜香属Ledum杜香L.palustre的果皮和种皮微形态特征进行观察,对杜鹃花属植物果皮微形态特征进行了系统描述,并通过比较现存杜鹃花属植物和种子化石,新发现一些居间的种子类型。结果表明,果皮和种皮微形态特征具有一定的系统学意义。叶状苞亚属subgen.Therorhodion的叶状苞杜鹃R.redowskianum的果实有短而疏的刺毛,无气孔,种子为无翅类,扁平,外围轮廓长椭圆形。杜鹃亚属subgen.Rhododendron植物果皮为百合花杜鹃型,具有鳞片,气孔器散生于指状突起之间,与叶表皮的微形态特征一致,其种子为百合花杜鹃型,表面具宽而浅的沟,呈脑纹状,有别于无鳞类杜鹃花。常绿杜鹃亚属subgen.Hymenanthes果皮为云锦杜鹃型,其角质层表面不规则,无表皮毛,偶见气孔;种子为云锦杜鹃型。映山红亚属subgen.Tsutsusi果皮为岭南杜鹃型,密生长表皮毛,角质层致密;种子为南边杜鹃型和岭南杜鹃型。微形态特征与“常绿杜鹃亚属和(国产)映山红亚属均为内部支持率很高的单系类群”的分子系统发育研究结果一致。马银花亚属subgen.Azaleastrum的马银花组sect.Azaleastrum和长蕊杜鹃组sect.Choniastrum微形态区别明显,支持各自为独立的单系类群。羊踯躅亚属subgen.Pentanthera的羊踯躅R.molle果皮特征明显,可与其他亚属区分,但种子类型更接近常绿杜鹃亚属。本研究结合分子系统发育资料和叶表皮微形态特征讨论了一些近缘类群的系统发育关系;研究结果支持将腺萼马银花处理为独立的种。     

An updated classification for the hyper-diverse genus Corydalis (Papaveraceae: Fumarioideae) based on phylogenomic and morphological evidence

The genus Corydalis, with ca. 530 species, has long been considered taxonomically challenging because of its great variability. Previous molecular analyses, based on a few molecular markers and incomplete taxonomic sampling, were clearly inadequate to delimit sections and subgenera. We have performed phylogenetic analyses of Corydalis and related taxa, using 65 shared protein-coding plastid genes from 313 accessions (including 280 samples of ca. 226 species of Corydalis) and 152 universal low-copy nuclear genes from 296 accessions (including 271 samples of Corydalis) covering all 42 previously recognized sections and five independent “series”. Phylogenetic trees were inferred using Bayesian Inference and Maximum Likelihood. Eight selected morphological characters were estimated using ancestral state reconstructions. Results include: (i) of the three subgenera of Corydalis, two are fully supported by both the plastid and nuclear data; the third, subg. Cremnocapnos, is weakly supported by plastid DNA only, whereas in the nuclear data the two included sections form successive outgroups to the rest of the genus; (ii) among all 42 sections and five “series”, 25 sections and one “series” are resolved as monophyletic in both data sets; (iii) the common ancestor of Corydalis is likely to be a perennial plant with a taproot, yellow flowers with a short saccate spur, linear fruits with recurved fruiting pedicels, and seeds with elaiosomes; (iv) we provide a new classification of Corydalis with four subgenera (of which subg. Bipapillatae is here newly described), 39 sections, 16 of which are consistent with the previous classification, 16 sections have been recircumscribed, one section has been reinstated and six new sections are established. Characters associated with lifespan, underground structures, floral spur, fruit and elaiosomes are important for the recognition of subgenera and sections. These new phylogenetic analyses combined with ancestral character reconstructions uncovered previously unrecognized relationships, and greatly improved our understanding of the evolution of the genus.     

Molecular and morphological phylogenetics of weevils (coleoptera,curculionoidea): do niche shifts accompany diversification?

The main goals of this study were to provide a robust phylogeny for the families of the superfamily Curculionoidea, to discover relationships and major natural groups within the family Curculionidae, and to clarify the evolution of larval habits and host-plant associations in weevils to analyze their role in weevil diversification. Phylogenetic relationships among the weevils (Curculionoidea) were inferred from analysis of nucleotide sequences of 18S ribosomal DNA (rDNA; approximately 2,000 bases) and 115 morphological characters of larval and adult stages. A worldwide sample of 100 species was compiled to maximize representation of weevil morphological and ecological diversity. All families and the main subfamilies of Curculionoidea were represented. The family Curculionidae sensu lato was represented by about 80 species in 30 "subfamilies" of traditional classifications. Phylogenetic reconstruction was accomplished by parsimony analysis of separate and combined molecular and morphological data matrices and Bayesian analysis of the molecular data; tree topology support was evaluated. Results of the combined analysis of 18S rDNA and morphological data indicate that monophyly of and relationships among each of the weevil families are well supported with the topology ((Nemonychidae, Anthribidae) (Belidae (Attelabidae (Caridae (Brentidae, Curculionidae))))). Within the clade Curculionidae sensu lato, the basal positions are occupied by mostly monocot-associated taxa with the primitive type of male genitalia followed by the Curculionidae sensu stricto, which is made up of groups with the derived type of male genitalia. High support values were found for the monophyly of some distinct curculionid groups such as Dryophthorinae (several tribes represented) and Platypodinae (Tesserocerini plus Platypodini), among others. However, the subfamilial relationships in Curculionidae are unresolved or weakly supported. The phylogeny estimate based on combined 18S rDNA and morphological data suggests that diversification in weevils was accompanied by niche shifts in host-plant associations and larval habits. Pronounced conservatism is evident in larval feeding habits, particularly in the host tissue consumed. Multiple shifts to use of angiosperms in Curculionoidea were identified, each time associated with increases in weevil diversity and subsequent shifts back to gymnosperms, particularly in the Curculionidae.     

Step-wise morphological trends in fluctuating environments: Evidence in the Late Devonian conodont genus Palmatolepis

Long-term morphological changes were investigated in the conodont genus Palmatolepis, using a Fourier analysis of the outline of platform elements. Three time-slices and four Late Frasnian - Early Famennian sections on North Gondwana were studied. No difference existed between locations, but shape variations were recorded associated with the successive Kellwasser events. The Middle Famennian period was analyzed in a section located at the Baltica margin across the annulata event. Some morphological changes may be related to the deposition of the corresponding black shales, but the main feature is a heterogeneity of the populations, which can be split into two morphological types. Latest Famennian conodonts from another North Gondwanan section show only minor shape changes during this time interval, which is environmentally stable prior to the global Hangenberg event. Based on synthetic shape variations across the whole period an morphological trend towards narrower elements is observed. This morphological change happens step-wise in response to global events. The amplitude of the shape shift is proportionally related to the extend of the successive Late Devonian global events.     

Molecules and morphology: Congruence of data inColumnea (Gesneriaceae)

The cladistic analysis and comparison of molecular and morphological data has been the source of much recent debate. In this study, independent analyses of molecular and morphological data fromColumnea L. sects.Pentadenia andStygnanthe (Gesneriaceae) are compared. Comparative methods include consensus, visually comparing trees from independent analyses and combined data analysis. Consensus methods provided little resolution. Comparison of trees obtained from the independent analyses revealed some differences although the trees are highly similar. However, a combined analysis found that the level of incongruence between the two data sets was low. The tree resulting from the combined data has aspects of both the morphological and molecular trees despite the larger number of molecular characters. In addition, the combined data tree has greater resolution than either of the two data sets singly, indicating that the two types of data are congruent, and complementary to each other.     

Tobacco to tomatoes: a phylogenetic perspective on fruit diversity in the Solanaceae

The Solanaceae contains many species of agricultural importance. Several of these are cultivated for their fruits, such as the tomato, the pepper and the aubergine. The family is very diverse in fruit type with capsules, drupes, pyrenes, berries, and several sorts of dehiscent non-capsular fruits occurring in the 90+ genera. In this paper, recent work on fruit type evolution in angiosperms is reviewed in relation to dispersal agents and habitat ecology. Defining fruit types in the Solanaceae in a simple five state system, then mapping them onto a previously published molecular phylogeny based on chloroplast DNA allows discussion of the evolution of these fruit types in a phylogenetic framework. Capsules are plesiomorphic in the family, and although berries are a synapomorphy (shared derived character) for a large clade including the genus Solanum (tomatoes and aubergines), they have arisen several times in the family as a whole. Problems with homology of drupes and pyrenes are discussed, and areas for future investigation of fruit structure homology identified. The distribution of fruit types in the large and diverse genus Solanum is also discussed in the light of monophyletic groups identified using chloroplast gene sequences. This variety is related to recent advances in the understanding of the molecular biology of fruit development. Finally, several key areas of future comparative, phylogenetic investigation into fruit type evolution in the family are highlighted.     

Otolith patterns of rockfishes from the northeastern pacific

Sagitta otolith shape was analysed in twenty sympatric rockfishes off the southern California coast (Northeastern Pacific). The variation in shape was quantified using canonical variate analysis based on fifth wavelet function decomposition of otolith contour. We selected wavelets because this representation allow the identifications of zones or single morphological points along the contour. The entire otoliths along with four subsections (anterior, ventral, posterodorsal, and anterodorsal) with morphological meaning were examined. Multivariate analyses (MANOVA) showed significant differences in the contours of whole otolith morphology and corresponding subsection among rockfishes. Four patterns were found: fusiform, oblong, and two types of elliptic. A redundancy analysis indicated that anterior and anterodorsal subsections contribute most to define the entire otolith shape. Complementarily, the eco‐morphological study indicated that the depth distribution and strategies for capture prey were correlated to otolith shape, especially with the anterodorsal zone. J. Morphol. 276:458–469, 2015. © 2014 Wiley Periodicals, Inc.     

QTL mapping of fruit-related traits in pepper (Capsicum annuum)

QTL analysis of pepper fruit characters was performed in an F₃ population derived from a cross between two Capsicum annuum genotypes, the bell-type cultivar Maor and the Indian small-fruited line Perennial. RFLP, AFLP^®1, RAPD and morphological markers (a total of 177) were used to construct a comparative pepper-tomato genetic map for this cross, and 14 quantitatively inherited traits were evaluated in 180 F₃ families. A total of 55 QTL were identified by interval analysis using LOD 3.0 as the threshold for QTL detection. QTL for several traits including fruit diameter and weight, pericarp thickness and pedicel diameter were often located in similar chromosomal regions, thus reflecting high genetic correlations among these traits. A major QTL that accounts for more than 60% of the phenotypic variation for fruit shape (ratio of fruit length to fruit diameter) was detected in chromosome 3. This chromosome also contained QTL for most of the traits scored in the population. Markers in linkage groups 2, 3, 8 and 10 were associated with QTL for multiple traits, thereby suggesting their importance as loci that control developmental processes in pepper. Several QTL in pepper appeared to correspond to positions in tomato for loci controlling the same traits, suggesting the hypothesis that these QTL may be orthologous in the two species.