Evolutionary transitions in parental care in cichlid fish

Cichlid fishes (Cichlidae) are well suited for testing theories of the evolution of vertebrate parental care. These freshwater teleost fish provide parental care for their offspring, display many different forms of care and have interspecific variation in which sex stays with the young. Here, we assemble the first family-wide composite phylogeny based on morphological and molecular studies, and trace two sets of character evolution: form of care (substrate guarding and mouthbrooding), and sex of care-giver (biparental, female-only, and male-only). Mouthbrooding has evolved from ancestral substrate guarding with 10 to 14 transitions and 0 to 3 reversals. The data support hypothesized transitions in the sex of care-giver, with uniparental female care having arisen from biparental care 21 to 30 times with 0 to 10 reversals. There is also evidence that male-only care evolved once from biparental care. These transitions in parental care characters are the most numerous reported for any family of vertebrates and, to our knowledge, provide the first quantitative support for models of parental care evolution in fish.     

A phylogenetic analysis of sex-specific evolution of ecological morphology in <Emphasis Type="Italic">Liolaemus</Emphasis> lizards

Adaptive radiation theory predicts that phenotypic traits involved in ecological performance evolve in different directions in populations subjected to divergent natural selection, resulting in the evolution of ecological diversity. This idea has largely been supported through comparative studies exploring relationships between ecological preferences and quantitative traits among different species. However, intersexual perspectives are often ignored. Indeed, although it is well established that intersexual competition and sex-specific parental and reproductive roles may often subject sex-linked phenotypes to antagonistic selection effects, most ecomorphological research has explored adaptive evolution on a single sex, or on means obtained from both sexes together. The few studies taking sexual differences into account reveal the occurrence of sex-specific ecomorphs in some clades of lizards, and conclude that the independent contribution of the sexes to the morphological diversity produced by adaptive radiation can be substantial. Here, we investigate whether microhabitat use results in the evolution of sex-specific ecomorphs across 44 Liolaemus lizard species. We found that microhabitat structure does not predict variation in body size and shape in either of the sexes. Yet, we found that males and females tend to occupy significantly different positions in multivariate morphological spaces, indicating that treating males and females as ecologically and phenotypically equivalent units may lead to incomplete or mistaken estimations of the diversity produced by adaptive evolution.     

Environmental and sibling resemblance components of variance and covariance in traits of early child development

The multiple regression analyses were undertaken to elucidate the significance and relative importance of different potential determinants of several child development traits (i.e. age at which child 1. turns himself over; 2. sits up; 3. stands up; 4. walks, and 5. cuts his first tooth). Despite the fact that we used a relatively wide gamut of potential determinants (such as parental age, geographic origin, occupation, current residence, or family size) of trait variability, the results of multiple regression analysis (N = 300 families) indicate an almost complete absence of significant factors and studied variables of development. Sibling resemblance component of variance and covariance for five mentioned traits of development and some morphological characters (weight, length and head circumference at day of birth and at 16 months) were studied in 66 families. The transmissibility ("heritability") values for the development traits were small, ranging from 0 ("Turn") to 0.42 ("Walked"). The Findings suggest that there are significant inverse relationship between some of the developmental traits and inborn morphological characters.     

Quantitative Ultrasound Characterization of Tumor Cell Death: Ultrasound-Stimulated Microbubbles for Radiation Enhancement

The aim of this study was to assess the efficacy of quantitative ultrasound imaging in characterizing cancer cell death caused by enhanced radiation treatments. This investigation focused on developing this ultrasound modality as an imaging-based non-invasive method that can be used to monitor therapeutic ultrasound and radiation effects. High-frequency (25 MHz) ultrasound was used to image tumor responses caused by ultrasound-stimulated microbubbles in combination with radiation. Human prostate xenografts grown in severe combined immunodeficiency (SCID) mice were treated using 8, 80, or 1000 µL/kg of microbubbles stimulated with ultrasound at 250, 570, or 750 kPa, and exposed to 0, 2, or 8 Gy of radiation. Tumors were imaged prior to treatment and 24 hours after treatment. Spectral analysis of images acquired from treated tumors revealed overall increases in ultrasound backscatter intensity and the spectral intercept parameter. The increase in backscatter intensity compared to the control ranged from 1.9±1.6 dB for the clinical imaging dose of microbubbles (8 µL/kg, 250 kPa, 2 Gy) to 7.0±4.1 dB for the most extreme treatment condition (1000 µL/kg, 750 kPa, 8 Gy). In parallel, in situ end-labelling (ISEL) staining, ceramide, and cyclophilin A staining demonstrated increases in cell death due to DNA fragmentation, ceramide-mediated apoptosis, and release of cyclophilin A as a result of cell membrane permeabilization, respectively. Quantitative ultrasound results indicated changes that paralleled increases in cell death observed from histology analyses supporting its use for non-invasive monitoring of cancer treatment outcomes.     

Phenotypic traits meet patterns of resource use in the radiation of “sharpfin” sailfin silverside fish in Lake Matano

Disruptive natural selection on traits related to resource exploitation may lead to differential adaptation and finally to speciation. Trait utility, the performance of traits in terms of fitness, is a central criterion for the recognition of adaptive radiation. Utility of morphological structures involved in foraging can be detected by relating their variation to individual resource use. Here, we test for trophic adaptations in the radiation of “sharpfin” sailfin silversides (Atheriniformes: Telmatherinidae), endemic to ancient graben-lake Matano in central Sulawesi (Indonesia). This small species-flock is characterized by high phenotypic diversity, including traits most likely related to feeding ecology. Previous analyses suggest that species boundaries are porous, indicating very recent or possibly ongoing processes of species flock formation. To test for adaptation to resource use in this radiation, we compared morphological traits among trophic groups of individuals as identified by stomach content analyses. We analyzed variation in candidate structures or structural complexes commonly recognized as indicative of trophic adaptation in fish radiations, including shapes of body, oral and pharyngeal jaws, gill rakers and body size. We found fine-scaled morphological differentiation according to feeding habits, covering all traits analyzed. Fish-, shrimp- and egg-feeders were most distinct, with major axes of morphological variation fitting patterns of adaptation reported from other lacustrine fish radiations. Thus, the present results are consistent with fine-scaled morphological adaptation to resource use, supporting the adaptive character of the sharpfin sailfin silverside radiation.     

Occurrence of stones, sediment and fish scales in stomach contents of demersal fish of the Rockall Trough

The occurrence of stones, sediment and fish scales was determined among the stomach contents of some 5000 fish. These fish comprised representatives of more than 70 different demersal species, sampled at 250 m bathymetric intervals, from the Rockall Trough, northeastern Atlantic Ocean, at depths ranging from 500 to 2900 m. Stones only occurred in stomachs of fish caught at 500–1000 m depth, their incidence in stomachs with food present being 4.6% at 500m, 1.1% at 750 m and 1.3% at 1000m depth. Sediment occurred in 9%, fish scales in 7%, of stomachs with contents. Sediment and scales co-occurred in the stomachs of primarily benthopelagic feeding fish. The sediment, however, was associated with the less dominant epibenthic components of the diets while the scales occurred with the more common benthopelagic components. The co-occurrence of sediment, scales and benthopelagic prey may arise through the former items having longer residence times in the stomachs than, for example, epibenthic prey. There is no evidence of increased occurrence of sediment (and its associated meiofauna) in the diets of fish on the lower slope at depths of 1500–2900 m in the Rockall Trough.     

The cost of secondary sexual characters and the evolution of cost-reducing traits

Secondary sexual traits are characterized by their exaggerated expression relative to homologous nonsexual characters in other species. All models of sexual selection assume that sex traits are costly to produce and maintain, and individuals with reduced costs of production and maintenance of secondary sexual characters would be at a selective advantage. A number of morphological, physiological and behavioural traits may have evolved as a result of their cost-reducing properties: (1) body size, which does not change throughout life, that allows certain individuals to develop exaggerated sex traits, (2) cost-reducing traits, such as muscle size, that improve with practice and (3) actual cost-reducing traits, such as wing size in birds with song flight, which are produced in advance of or simultaneously with the sex trait. Cost-reducing traits may coevolve with secondary sexual characters and allow more extreme sexual signalling than would otherwise have been possible in their absence or in reduced versions.     

The cost of secondary sexual characters and the evolution of cost-reducing traits

Secondary sexual traits are characterized by their exaggerated expression relative to homologous nonsexual characters in other species. All models of sexual selection assume that sex traits are costly to produce and maintain, and individuals with reduced costs of production and maintenance of secondary sexual characters would be at a selective advantage. A number of morphological, physiological and behavioural traits may have evolved as a result of their cost-reducing properties: (1) body size, which does not change throughout life, that allows certain individuals to develop exaggerated sex traits, (2) cost-reducing traits, such as muscle size, that improve with practice and (3) actual cost-reducing traits, such as wing size in birds with song flight, which are produced in advance of or simultaneously with the sex trait. Cost-reducing traits may coevolve with secondary sexual characters and allow more extreme sexual signalling than would otherwise have been possible in their absence or in reduced versions.     

蜈蚣草耐铅、铜、锌毒性和修复能力的研究

重金属复合污染是主要土壤污染类型之一。为了探明蜈蚣草修复土壤重金属复合污染的能力,土培试验研究了分别添加不同浓度铅、铜、锌条件下蜈蚣草生物量变化,结果表明,不同浓度铅和较低浓度锌处理下蜈蚣草生物量呈显著性地增加,土壤Pb、Zn添加浓度分别为1750mg／kg和1350mg／kg,蜈蚣草生物量最大。说明蜈蚣草有极强的耐Pb、Zn毒性能力,能在较高有效态Pb或Zn污染土壤上正常生长。蜈蚣草具有一定的耐Cu毒性能力,在Cu耐性方面,蜈蚣草可能存在生态型的差异。以上结果说明,利用蜈蚣草修复萃取Pb—As,Zn—As,Cu-As等复合污染土壤上As有重要的意义。     

Transgressive character expression in a hybrid sunflower species

Diploid hybrid lineages often are ecologically distinct from their parental species. However, it is unclear whether this niche divergence is typically achieved via hybrid intermediacy, a mixture of parental traits, and/or the evolution of extreme (transgressive) morphological and ecophysiological features. Here we compare an extensively studied hybrid sunflower species, Helianthus anomalus, with its putative parents, H. annuus and H. petiolaris, for 41 morphological and 12 ecophysiological traits. Helianthus anomalus was morphologically intermediate for one trait (2.4%), parental-like for 23 traits (56.1%), and transgressive for 17 traits (41.5%). For ecophysiological traits, H. anomalus was not significantly different from one or both parents for nine traits (75%), and was transgressive for the remaining three (25%). Thus, H. anomalus appears to be a mosaic of parental-like and transgressive phenotypes. Although the fitness effects of the transgressive characters are not yet known, many of these characters are consistent with adaptations reported for other sand dune plants. Genetic studies are currently underway to ascertain whether these extreme characters arose as a direct byproduct of hybridization or whether they evolved via mutational divergence.     

Character analysis in morphological phylogenetics: problems and solutions

Many aspects of morphological phylogenetics are controversial in the theoretical systematics literature and yet are often poorly explained and justified in empirical studies. In this paper, I argue that most morphological characters describe variation that is fundamentally quantitative, regardless of whether they are coded qualitatively or quantitatively by systematists. Given this view, three fundamental problems in morphological character analysis (definition, delimitation, and ordering of character states) may have a common solution: coding morphological characters as continuous quantitative traits. A new parsimony method (step-matrix gap-weighting, a modification of Thiele's approach) is proposed that allows quantitative traits to be analyzed as continuous variables. The problem of scaling or weighting quantitative characters relative to qualitative characters (and to each other) is reviewed, and three possible solutions are described. The new coding method is applied to data from hoplocercid lizards, and the results show the sensitivity of phylogenetic conclusions to different scaling methods. Although some authors reject the use of continuous, overlapping, quantitative characters in phylogenetic analysis, quantitative data from hoplocercid lizards that are coded using the new approach contain significant phylogenetic structure and exhibit levels of homoplasy similar to those seen in data that are coded qualitatively.     

Immune responses,intestinal microbiota,performance and blood characteristics of Japanese quail fed on diets containing camphor

This study aimed to investigate the effect of supplemental camphor on the performance and immune functions of Japanese quail by feeding graded levels (0 (control), 250, 500, 750, 1000, 5000 or 10 000 ppm) of camphor during a 42-day feeding trial. In all, 280 1-day-old quail chicks were randomly assigned into 28 cages of 10 chicks each with separate feeders. The results clearly demonstrated that camphor did not have a significant effect on BW, BW gain, total experimental average daily feed intake, feed conversion ratio, internal organ relative weights and biochemical parameters such as uric acid, albumin, total protein and triglyceride; however, plasma cholesterol concentration was significantly different in a linear manner, in which 500 ppm of camphor resulted in a lower level of cholesterol. Alternatively, greater concentrations of glucose and high-density lipoprotein (HDL) were also found in 5000 and 1000 ppm of camphor groups, respectively. Cellular responses to the phytohaemagglutinin-P and 2,4-dinitro 1-chlorobenzene skin test were not influenced by dietary camphor. Humoral responses to secondary sheep red blood cells, avian influenza virus (AIV) and Newcastle disease virus (NDV) immunisations were positively influenced by camphor supplementation, in which greater secondary response to sheep erythrocytes belonged to 750 and 1000 ppm of camphor groups; whereas, diet supplementation with camphor had no significant effect on lymphoid organ weights and heterophil-to-lymphocyte ratio. The greatest AIV antibody titers were seen in groups, which received 1000 and 5000 ppm of camphor (P<0.05) and the values of NDV antibody titers increased with an increase in the camphor consumption. Furthermore, dietary inclusion of 500 ppm of camphor positively decreased coliform populations in the gastrointestinal tract (GIT). In addition, an increase in lactic acid bacteria was also observed in quails, which were fed on the diets containing 750 ppm camphor. Collectively, these data suggest that as a phytogenic feed additive, camphor may effectively act as a modulator of health status (by increasing glucose and HDL), GIT microbiota and immunological responses of the Japanese quail.     

BOTH MALE AND FEMALE SEXUAL ORNAMENTS REFLECT OFFSPRING PERFORMANCE IN A FISH

Secondary sexual characters are often expressed in both sexes (mutual ornamentation), but are less often studied simultaneously. We studied the adaptive signaling function of male and female ornamentation in a mutually ornamented fish, the whitefish Coregonus lavaretus. In an experimental design in which nongenetic environmental effects were minimized, we found that highly ornamented females, males, and their parental combinations had offspring with better swimming performance and predator‐avoidance ability than less ornamented individuals or combinations. Furthermore, highly ornamented females had larger offspring that also had higher yolk volume than less ornamented individuals. Offspring swimming performance was not dependent on offspring size and was only weakly affected by yolk volume, which suggest that swimming performance and measured morphological traits are independent fitness measures. In conclusion, mutual ornamentation of whitefish may signal the quality of individuals in both sexes, which may indicate ongoing directional selection for these ornamental traits. However, offspring fitness traits were also dependent on parental combination, which suggests that genetic compatibility effects may weaken the directional selection and the indicator value of the ornamentation.     

The effect of isozyme selection on metric characters in an interspecific backcross of tomato — basis of an early screening procedure

Summary The extent of correlation was estimated between isozyme genotypes and the four widely segregating characters — leaf segment W/L ratio, stigma exsertion, fruit weight, and seed weight — in the first backcross of F₁ Lycopersicon esculentum x Solanum pennellii to the former parent. The inbred parents differ in their alleles at the 12 tested isozymic loci, which are known to mark a minimum of eight of the twelve tomato chromosomes. Based on the isozyme data, a mean heterozygosity value, ¯H, was calculated which estimates the proportion of pennillii alleles in each individual. Correlations between mean heterozygosity and observed levels of each quantitative trait were highly significant and positive or negative as expected from the relative parental values. Plants with the lowest mean heterozygosity — i.e., closest to the esculentum zymotype also had mean values closest to those of this parent amongst the whole backcross population for each of the quantitative traits.Bivariate and multiple regression analysis was used to evaluate the ability of isozymes vs diagnostic morphological characters to estimate the portion of recurrent parent genes carried in each backcross individual. The results suggest that isozyme data gives better estimates than single diagnostic morphological characters and approach the level obtained by combinations of three morphological traits. Since electrophoretic determinations are made on small seedlings, selection at that stage can effect great savings of space and effort by greatly deminishing the size of the population needed at maturity. As such, isozyme selection would precede morphological selection but not replace it, thus the predictive value of these biochemical markers as well as diagnostic morphological characters could be obtained.     

Natural hybridization and hybrid zones between Quercus crassifolia and Quercus crassipes (Fagaceae) in Mexico: morphological and molecular evidence

Hybrid zones provide interesting systems to study genetic and ecological interaction between different species. The correct identification of hybrids is necessary to understand the evolutionary process involved in hybridization. An oak species complex occurring in Mexico formed by two parental species, Quercus crassifolia H. & B. and Q. crassipes H. & B., and their putative hybrid species, Q. dysophylla, was analyzed with molecular markers (random amplified polymorphic DNA [RAPDs]) and morphological tools in seven hybrid zones (10 trees per taxa in each hybrid zone) and two pure sites for each parental species (20 trees per site). We tested whether geographic proximity of hybrid plants to the allopatric site of a parental species increases its morphological and genetic similarity with its parent. Seventeen morphological traits were measured in 8700 leaves from 290 trees. Total DNA of 250 individuals was analyzed with six diagnostic RAPD primers. Quercus crassifolia differed significantly from Q. crassipes in all the examined characters. Molecular markers and morphological characters were highly coincident and support the hypothesis of hybridization in this complex, although both species remain distinct in mixed stands. Clusters and a hybrid index (for molecular and morphological data) showed that individuals from the same parental species were more similar among themselves than to putative hybrids, indicating occasional hybridization with segregation in hybrid types or backcrossing to parents. Evidence does not indicate a unidirectional pattern of gene flow.     

The effects of age,sex, and habitat on body size and shape of the blackstripe topminnow,Fundulus notatus (Cyprinodontiformes: Fundulidae) (Rafinesque 1820)

Lake and stream habitats pose a variety of challenges to fishes due to differences in variables such as water velocity, habitat structure, prey community, and predator community. These differences can cause divergent selection on body size and/or shape. Here, we measured sex, age, length, and eight different morphological traits of the blackstripe topminnow, Fundulus notatus, from 19 lake and stream populations across four river drainages in central Illinois. Our goal was to determine whether size and shape differed consistently between lake and stream habitats across drainages. We also considered the effects of age and sex as they may affect size and morphology. We found large differences in body size of age 1 topminnows where stream fish were generally larger than lake fish. Body shape mainly varied as a function of sex. Adult male topminnows had larger morphological traits (with the exception of body width) than females, in particular longer dorsal and anal base lengths. Subtle effects of habitat were present. Stream fish had a longer dorsal fin base than lake fish. These phenotypic patterns may be the result of genetic and/or environmental variation. As these lakes are human‐made, the observed differences, if genetic, would have had to occur relatively rapidly (within about 100 years). © 2013 The Linnean Society of London     

Characteristics of the submicroscopic structure of hybrid clone cells obtained by the fusion of mouse hepatoma cells and mink fibroblasts and containing different numbers of mink chromosomes

Four hybrid clones obtained by fusion of mouse hepatoma cells with mink fibroblasts treated with polyethyleneglycol were studied morphologically and morphometrically using electron microscopy. The clones studied contained a double set of mouse chromosomes and different numbers of mink chromosomes. It is demonstrated that clones containing different mink chromosomes differ considerably from each other and from the parental cells in the manifestation of some morphological characters (form and type of cell growth, form of the nucleus, structure of mitochondria, distribution of membranes of the granular endoplasmic reticulum), as well as in some quantitative parametres of organelles (area of the cut of the cell and of the nucleus, a relative volume of the nucleus). The data obtained witness for the fact that some morphological traits characteristic of cells of a certain parental type may appear in hybrids independently of each other, and that the degree of their manifestation may depend on the number of chromosomes of one of the parents or, possibly, on one particular chromosome.     

Hybridization and transgressive exploration of colour pattern and wing morphology in Heliconius butterflies

Hybridization can generate novel phenotypes distinct from those of parental lineages, a phenomenon known as transgressive trait variation. Transgressive phenotypes might negatively or positively affect hybrid fitness, and increase available variation. Closely related species of Heliconius butterflies regularly produce hybrids in nature, and hybridization is thought to play a role in the diversification of novel wing colour patterns despite strong stabilizing selection due to interspecific mimicry. Here, we studied wing phenotypes in first‐ and second‐generation hybrids produced by controlled crosses between either two co‐mimetic species of Heliconius or between two nonmimetic species. We quantified wing size, shape and colour pattern variation and asked whether hybrids displayed transgressive wing phenotypes. Discrete traits underlain by major‐effect loci, such as the presence or absence of colour patches, generate novel phenotypes. For quantitative traits, such as wing shape or subtle colour pattern characters, hybrids only exceed the parental range in specific dimensions of the morphological space. Overall, our study addresses some of the challenges in defining and measuring phenotypic transgression for multivariate traits and our data suggest that the extent to which transgressive trait variation in hybrids contributes to phenotypic diversity depends on the complexity and the genetic architecture of the traits.     

Leaf epidermal features of Rhododendron (Ericaceae)from China and their systematic significance

Rhododendron is the largest genus within the subfamily Rhododendroideae, which has about 1000 known species in the world and more than 500 species in China. Since the genus was established by Linnaeus, its infrageneric relationships have been well studied by many taxonomists on the basis of morphological characters and molecular data. In 1996, Chamberlain et al. proposed a new system of Rhododendron with eight subgenera, i.e., Azaleastrum, Candidastrum, Hymenanthes, Mumeazalea, Pentanthera, Rhododendron, Therorhodion, and Tsutsusi. In this paper, micromorphological characters of leaf epidermis in 4 varieties, 48 species, 6 subgenera of Rhododendron from China were examined using light microscopy (LM) and scanning electron microscopy (SEM). Leaf epidermal features are described and micromorphological types are distinguished here according to morphological characters such as scale, gland, foliar trichome and stomatal apparatus of leaf epidermis. It is shown that the leaf epidermal cells are usually irregular or polygonal in shape. The patterns of anticlinal walls are straight, arched or undulate. The stomatal apparatuses are anomocytic and are usually found on abaxial, not adaxial, epidermis. The results also show that: (1) the lepidote rhododendron (i.e., subgen. Rhododendron), which has both scales and papillae on leaf epidermis, differs distinctly from the elepidote rhododendron; (2) three types of leaf epidermis are identified in subgen. Hymenanthes (i.e., R. fortunei-type, R. chihsinianum-type and R. simiarum-type), whereas four in subgen. Tsutsusi (i.e., R. mariesii-type, R. simsii-type, R. mariae-type and R. flosculum-type); (3) except for R. westlandii and R. henryi, the species of subgen. Azaleastrum show similar morphological characters, i.e., dense stomatal apparatuses surrounded by ringed or discontinuous striates; (4) R. molle of subgen. Pentanthera differs from the species of other subgenera on morphological characters such as foliar trichomes, dense stomatal apparatuses with asymmetrical outer stomatal rims surrounded by undulate-striates, and no gland; (5) only R. redowskianum is found with distinct T-pieces at the polar region of guard cells in Rhododendron. The results support the conclusion inferred from molecular systematic studies that subgen. Therorhodion is the basal clade of Rhododendron. Finally, the relationships between the closely related species are also discussed on the basis of leaf epidermal features.     

中国杜鹃花属的叶表皮特征及其系统学意义

通过光学显微镜和扫描电镜对国产杜鹃花属Rhododendron6个亚属48种4变种植物的叶表皮特征进行观察,结果表明叶表皮的鳞片、腺体、表皮毛的有无和气孔器的类型具有一定的系统学意义。光学显微镜下可见：叶表皮细胞形状为多边形、不规则形,垂周壁式样为较平直、浅波状和波状．气孔器普遍存在于下表皮,上表皮未见气孔器分布,其类型均为无规则形。杜鹃亚属subgen．Rhododendron叶表皮具有鳞片,在扫描电镜下可见气孔器散生于指状突起之间,有别于无鳞类杜鹃花。常绿杜鹃亚属subgen．Hymenanthes和映山红亚属subgen．Tsutsusi有多种不同叶表皮类型;除了凯里杜鹃R．westlandii和弯蒴杜鹃R．henryi形态较特殊之外,马银花亚属subgen．Azaleastrum的马银花组sect．Azaleastrum和长蕊杜鹃组sect．Choniastrum微形态区别不明显。羊踯躅亚属subgen．Pentanthera的羊踯躅R．molle叶表皮有表皮毛,无腺体,气孔器密生,外拱盖形状不对称,外围有多层波形条纹突起可与其他亚属区分：而叶状苞亚属subgen．Therorhodion的叶状苞杜鹃R．redowskianum保卫细胞有明显的T型加厚,与其他亚属种类明显不同,与分子系统发育研究得到的“叶状苞亚属是杜鹃花属的基部类群”的结果吻合。本研究还依据叶表皮微形态特征讨论了一些近缘种类的关系。