White Color Morphs Research Articles

Herbivore pressure by weevils associated with flower color polymorphism in Geranium thunbergii (Geraniaceae)

Although floral herbivory has recently received increased attention as an important factor influencing plant reproduction, relatively little is known about how its frequency and intensity vary depending on traits of host plants. Here we report that herbivore pressure by a weevil, Zacladus geranii, is associated with a flower color polymorphism of Geranium thunbergii (Geraniaceae). Pink and white flower color morphs have been reported in G. thunbergii, and we found in a three-year field survey in multiple populations that, generally, adult weevils more preferentially visited white flowers than pink flowers. Consistently, we found more severe damage by weevil larvae in white flowers. Overall herbivore pressure for G. thunbergii varied strongly between populations, and the difference seems to be partly explained by the co-occurrence of a related plant species, Geranium yezoense, in a population, as weevils preferred it to both color morphs of G. thunbergii, thereby relaxing overall herbivore pressure for G. thunbergii. Nonetheless, despite such high variability, the preference of weevils for white morphs over pink morphs of G. thunbergii was found across multiple populations. We discuss possible mechanisms causing the association between flower color and herbivore preference as well as its evolutionary consequences.

Genetic diversity and population structure of North America’s rarest heron, the reddish egret (Egretta rufescens)

The global distribution of the reddish egret is characterized by disjunct colonies occurring from the Pacific side of Northwest Mexico to the Caribbean. We examined distantly isolated colonies of reddish egret to determine global population genetic structure. We used seven polymorphic microsatellites to accomplish five goals: (1) to assess range wide population differentiation among reddish egret (Egretta rufescens) populations, (2) identify extent of gene flow, (3) determine any historical occurrence of bottlenecks, (4) assess genetic differentiation between color morphs, (5) clarify subspecies status of E. r. dickeyi, a completely dark morph population located in and around the Baja California peninsula, Mexico. Genetic differentiation was dramatic (global Fst = 0.161) throughout the reddish egrets range extending from Baja California, Mexico to Great Inagua, Bahamas. Differentiation occurred among three distinct regions (Fst = 0.238) but not among colonies/islands within regions suggesting regional philopatry. Genetic diversity (alleles per locus, and heterozygosity) in Baja California Sur, Mexico and Great Inagua, Bahamas populations is lower than in the Texas/Mexico population due to minimal dispersal between regions and smaller population sizes. Dark and white color morphs when present within the same region showed no differentiation. Patterns of recent population bottlenecks are not evident in each of the three regional populations. With evidence of limited gene flow in addition to low genetic diversity and prospects of habitat loss we recommend that reddish egrets be managed as three distinct or evolutionary significant units.

The Impact of Biochemistry vs. Population Membership on Floral Scent Profiles in Colour Polymorphic Hesperis matronalis

Studies of floral scent evolution often attribute variation in floral scent to differences in pollinator behaviour, ignoring the potential for shared biochemistry between floral scent and floral colour to dictate patterns of phenotypic variation in scent production. To determine the relative effects of shared biochemistry and/or localized population-level phenomena on floral scent phenotype, floral scent composition and emission rate were examined in five wild populations of colour polymorphic Hesperis matronalis (Brassicaceae). Floral scent was collected by in situ dynamic headspace extraction on purple and white colour morphs in each of five wild populations. Gas chromatography-mass spectroscopy of extracts allowed determination of floral scent composition and emission rate for all individuals, which were examined by non-metric multidimensional scaling and analysis of variance (ANOVA), respectively, to determine the contributions of floral colour and population membership to scent profile variation. Despite the fact that colour morph means were very similar in some populations and quite different in other populations, colour morphs within populations did not differ from each other in terms of scent composition or emission rate. Populations differed significantly from one another in terms of both floral scent composition and emission rate. Shared biochemistry alone cannot explain the variation in floral scent phenotype found for H. matronalis. Such a result may suggest that the biochemical association between floral scent and floral colour is complex or dependent on genetic background. Floral scent does vary significantly with population membership; several factors, including environmental conditions, founder effects and genetics, may account for this differentiation and should be considered in future studies.

Comparative life history and parasitism of a new colour morph of the walnut aphid in California

1 The walnut aphid Chromaphis juglandicola is a yellow aphid. In 2003, however, a white colour morph was discovered in the Sacramento Valley of California. The colour dimorphism occurs between clone lines and, when white morphs are present, they occur in mixed colour morph colonies on the underside of walnut leaves. 2 Laboratory experiments were undertaken to evaluate the thermal requirements for development, adult longevity and progeny production of the two colour morphs. Host instar preference of Trioxys pallidus, a parasitoid responsible for the successful biological control of the walnut aphid in California, was examined separately for each colour morph, and host colour preference was investigated for the preferred instar. 3 No differences in thermal requirements for development, adult size or mean longevity were detected between yellow and white colour morphs. A small difference in early reproduction was detected: white colour morphs produced more progeny on each of the two first days of adult reproduction than yellow colour morphs. 4 Trioxys pallidus showed a slight preference for the fourth instar of the yellow morph over the second- and third-, but equal preference for second, third and fourth instars of the white morph. When offered equal numbers of fourth instars of the two colour morphs, T. pallidus did not show any colour preference. 5 The differences in early aphid reproduction and host instar preference by T. pallidus were combined in a stage-structured matrix model. Model analysis showed a greater potential for population growth of the white morph over the yellow morph, with early reproduction having a greater influence than host instar preference.

Flower color–flower scent associations in polymorphic Hesperis matronalis (Brassicaceae)

Floral scent emission rate and composition of purple and white flower color morphs of Hesperis matronalis (Brassicaceae) were determined for two populations and, for each, at two times of day using dynamic headspace collection and GC–MS. The floral volatile compounds identified for this species fell into two main categories, terpenoids and aromatics. Principal component analysis of 30 compounds demonstrated that both color morphs emitted more scent at dusk than at dawn. Color morphs varied in chemical composition of scent, but this differed between populations. The white morphs exhibited significant differences between populations, while the purple morphs did not. In the white morphs, one population contains color–scent associations that match expectations from classical pollination syndrome theory, where the flowers have aromatic scents, which are expected to maximize night-flying moth pollinator attraction; in the second population, white morphs were strongly associated with terpenoid compounds. The potential impact that pollinators, conserved biosynthetic pathways, and the genetics of small colonizing populations may have in determining population-specific associations between floral color and floral scent are discussed.

Systematic significance of tridentate pedicellariae in the echinoid genera Diadema and Echinothrix

Abstract. The tridentate pedicellariae in all species of Diadema and Echinothrix recognized by Mortensen, plus species described later (to include D. palmeri and two color morphs of E. calamaris) were examined to identify diagnostic species characters, and to determine whether such characters are useful in inferring phylogenetic relationships between species. Nineteen morphological characters were measured and analyzed, and species‐specific characters were identified. The morphometric data were then transformed using gap coding and a parsimony analysis was undertaken. The resulting cladogram for the tridentate pedicellariae present in species of Diadema was compared with the mitochondrial DNA phylogeny of Lessios et al., with a good level of congruence observed. Very narrow forceps‐like tridentate pedicellariae were found to be basal among Recent species, present only in D. setosum and D. palmeri. Members of D. palmeri were found to be unique within the genus, having both the forceps‐like form and a broad form of tridentate pedicellaria. The very narrow forceps‐like form is absent in all subsequent divergent species, while the broad form of tridentate pedicellaria in D. palmeri is basal to the broad forms in D. antillarum, D. mexicanum, D. paucispinum, and both the broad and narrow forms of tridentate pedicellaria of D. savignyi. The greatest similarity was found between the broad forms of tridentate pedicellaria in D. antillarum and D. mexicanum. These species were also found to have narrow forms of tridentate pedicellaria that showed a reasonable level of association to one another and to the rostrate form in D. ascensionis. Since the work of Mortensen, D. ascensionis has been shown, using mitochondrial DNA, to be nested within D. antillarum. The results of this study show that the tridentate pedicellaria in D. ascensionis are distinctly different from all other forms within the genus, to the extent that they belong to a rostrate subclass. This indicates that, among Diadema species, the tridentate pedicellariae are one of the first morphological characters to diverge in genetically isolated populations. Unlike the tridentate pedicellariae in Diadema, the different forms of tridentate pedicellariae in E. calamaris and E. diadema showed a stronger association intraspecifically than between forms interspecifically. The small forms of tridentate pedicellaria of the white and brown color morphs in E. calamaris showed a high level of association, while the large forms showed only a moderate association, far lower than would be expected between color morphs of the same species. This suggests that they are either subspecies or recently diverged separate species. This study demonstrates the taxonomic value contained within the characters of the tridentate pedicellariae for both differentiating species (even subspecies) and inferring phylogenetic relationships between species.

Lunar periodicities of diadematid echinoids breeding in Fiji

Gonad indices for two species of Diadema and two species of Echinothrix, including two color morphs of Echinothrix calamaris, are described for a 12-month period on Sosoikula Reef and Nukubuco Reef, Viti Levu, Fiji. Seasonal fluctuations in salinity (36.11±0.88 ppt) and water temperature (26.35±0.91°C) occurred. Gonad index data showed monthly reproductive rhythms, closely attuned to the lunar cycle. Diadema savignyi and Echinothrix diadema spawned on the full moon, and Diadema setosum and E. calamaris (white and brown color morphs) spawned on the new moon. Breeding periodicities coincided with the spring tides, thus maximizing chances of fertilization and dispersal. Such breeding cycles indicate how closely related species can co-exist with minimum risk of hybridization. Unusual sex ratios were found for all species, with an exceptionally low incidence of males. Reasons for this deviation from the typically reported 1:1 sex ratio may relate to the exceptionally high levels of tributyltin (TBT) recorded in Suva Harbor.

Taxonomic significance of spine morphology in the echinoid genera Diadema and Echinothrix

Abstract. The spine morphology of all established species of Diadema and Echinothrix, including 2 color morphs of E. calamaris, were examined externally and internally via transverse sectioning to identify diagnostic species features and to assess the morphological relationship between species. Forty‐nine different morphological characters were measured and analysed using ordination by multi‐dimensional scaling (MDS) and cluster analysis. Specimens of Diadema paucispinum and D. setosum had very distinct spine structures. In D. paucispinum, the spines were more robust than those of other species of Diadema. This was evident in the spine's internal structure, with large, closely packed solid wedges, a small axial cavity, and rings of trabeculae throughout the spine's length. The spines in D. setosum were distinctive because of their length in relation to test size and the reduced flaring of their verticillations. The spines of other members of this genus were very similar to each other. Without careful sectioning, the spines from specimens of D. antillarum, D. ascensionis, D. mexicanum and D. savignyi were difficult to differentiate. The internal structures of spines for each species did, however, possess a combination of features that differentiated the species. Such features included the shape, orientation, and number of solid wedges, the presence or absence of spokes and rings of trabeculae between the solid wedges, and the presence or absence of tissue within the axial cavity. Individuals of Diadema palmeri also had spines morphologically similar to other species, however, the red pigmentation of these spines (in life and when preserved) made them easily distinguishable. The spine structures of the 2 species of Echinothrix were starkly different, while the white and brown color morphs of E. calamaris had morphologically distinctive ambulacral and interambulacral spines. The blunt, open‐tipped interambulacral spines, with reticular tissue present in the axial cavity of the white color morph, were easily distinguished from the pointed, closed‐tipped spines, with a hollow axial cavity found in the brown color morph. Such differences indicate that the brown color morph is either a subspecies or a separate species. Taken together the data show that each species has significant morphological differences in the structure of the spines. It is evident from our data that spine morphology is a useful tool to differentiate these commonly confused species.

Tan and White Color Morphs of White-Throated Sparrows Differ in Their Non-Song Vocal Responses to Territorial Intrusion

White-throated Sparrows (Zonotrichia albicollis) occur in two distinct color morphs indicated by the color of the bird's median crown stripe. Tan-striped (TS) and white-striped (WS) sparrows differ in their aggressive behavior, with WS males generally behaving more aggressively than TS males. We measured the number and type of songs and call vocalizations used by TS and WS males in response to simulated territorial intrusion. TS males used significantly more low aggression calls compared to WS males. These results suggest that non-song call vocalizations, in addition to measures of song, may be valuable for future studies of territorial behavior and communication in birds.

Foliar Herbivory Affects Floral Characters and Plant Attractiveness to Pollinators: Implications for Male and Female Plant Fitness

Foliar Herbivory Affects Floral Characters and Plant Attractiveness to Pollinators: Implications for Male and Female Plant Fitness

Population fluctuations of gyrfalcon and rock ptarmigan: analysis of export figures from Iceland

We analysed harvest data for gyrfalcon Falco rusticolus and rock ptarmigan Lagopus mutus from Iceland with respect to regularity in fluctuations of numbers. The gyrfalcon data concerned live trapped birds exported to Denmark between 1731 and 1793, and totalled 4,848 falcons, including 4,318 grey, 156 half‐white and 374 white colour morphs. According to contemporary sources grey birds were part of the local breeding population (islandus‐type birds) but the other morphs represented mainly visitors from Greenland. This is also the current situation but some of the lightest Icelandic breeders could be classified as half‐white. The rock ptarmigan harvest data concerned birds exported to Europe in the period 1864–1919, in total ca 3.3 million birds. The data series for white and half‐white gyrfalcons were significantly correlated (r = 0.501, p < 0.001). The data series for grey and white morphs (r = ‐0.099, P = 0.445) and grey and the half‐white morphs (r = ‐0.1183, P = 0.360), showed no correlation. Time series analysis showed that the white (candicans‐type) morph fluctuated irregularly. The half‐white morph behaved similarly but also showed some affinity with the grey morph, and could have represented a mixture of local breeders and Greenlandic winter visitors. Grey morph gyrfalcons and rock ptarmigan showed regular fluctuations in numbers with a 10‐year periodicity. The reliance of Icelandic gyrfalcons on rock ptarmigan during the early part of the breeding season and in all phases of the ptarmigan cycle is well established and may offer a case for causal connections between the two cyclic populations.

POLLINATION AND SEED PRODUCTION IN IPOMOPSIS AGGREGATA: DIFFERENCES AMONG AND WITHIN FLOWER COLOR MORPHS

The pollination of red, pink, and white color morphs of Ipomopsis aggregata was evaluated to assess whether ethological isolation based on pollinator color discrimination may occur. We observed animal visitors, assessed pollen delivery, seed set per fruit, percentage of flowers setting fruit, nectar production, and timing of flower opening for different color morphs in the Front Range of Colorado. Based on traditional zoophilous flower classifications, we expected hummingbirds to pollinate red‐flowered I. aggregata subsp. collina and hawkmoths to pollinate white‐flowered I. aggregata subsp. Candida. However, ethological isolation does not appear to occur among color morphs of I. aggregata in the Front Range. Hummingbirds visited red‐flowered plants in excess overall, and, to a lesser extent, so did hawkmoths. Both hummingbirds and hawkmoths visited all color morphs and probably transferred pollen among them. Pollen delivery data and a day‐night bagging experiment also suggest that pollinators do not necessarily behave as predicted by flower classifications. In addition, there is little evidence for major differences between red, white, and pink flowers in any aspects of reproductive biology. Indeed, most variation occurs within a given color morph.

The Morphological Identification of Domesticated Zebra Finches, Poephila-Guttata (Passeriformes, Estrilidae), in Australia

The zebra finch, Poephila guttata, is a native Australian grassfinch which is used extensively in laboratories of many countries, including Australia. We have compared the morphology of groups of zebra finches from various origins: (1) wild-caught from a population at Bunbartha in northern Victoria; (2) their first-generation offspring which were bred in captivity; (3) fawn colour morphs; (4) white colour morphs; (5) a private collection which included a variety of colour morphs. Birds in groups 3,4 and 5 were considered to be the distant offspring of wild-caught zebra finches. The length of the wing, the head and the tarsus, the width and the depth of the bill, and the weight were measured. All characters, except the width of the bill, differed significantly among the groups. Analyses of variance indicated that head length, tarsus length and bill depth differed significantly between group 1 and groups 3, 4 and 5, but were not significantly different between groups 1 and 2 nor among groups 3,4 and 5. The wild-caught birds and their immediate offspring were larger than those of the colour-morph groups. Discriminant analysis showed that wing length, tarsus length and bill depth were important in classifying individuals into groups. Differences between groups were attributed to the effects of domestication upon birds in aviaries.