Morph Type Research Articles

Impact of mate availability, population size, and spatial aggregation of morphs on sexual reproduction in a distylous, aquatic plant

In distylous, self-incompatible plants, clonal propagation, unbalanced floral morph frequencies, and reduced population size can interfere with the functioning of distyly by compromising legitimate intermorph pollinations, resulting in reduced reproductive output. Here, we examined the mating system and the impact of mate availability, population size, and spatial aggregation of morphs on reproductive output in the distylous, clonal, aquatic plant Hottonia palustris. Controlled pollinations under greenhouse conditions detected no spontaneous selfing without the action of a pollen vector (autonomous autogamy) and demonstrated very low fruit and seed development after self-pollination. Intermorph (legitimate) crossings resulted in high reproductive output in both floral morphs (long- and short-styled individuals), whereas intramorph (illegitimate) crossings decreased fruit and seed development by more than 50%, indicating that the species has partial intramorph-incompatibility. In natural populations, small population size and increasing deviation of floral morph frequencies negatively affected reproductive outcome. Individuals of the majority morph type developed significantly fewer fruit and seeds than individuals of the minority morph type. This rapid decline in fecundity was symmetrical, indicating that regardless of which morph was in the majority, the same patterns of negative frequency-dependent mating occurred. Increasing spatial isolation between compatible morphs significantly reduced fruit and seed set in both morphs similarly. This study provides clear indications of frequency- and context-dependent mating in natural populations of a distylous plant species.

Morph-specific differences in reproductive success in the distylous Primula veris in a context of habitat fragmentation

Abstract Heterostylous self-incompatible plant species are particularly sensitive to habitat fragmentation and to disruption of pollination processes because of the need of intermorph cross-pollination for producing seeds. Heterostyly is characterized by sexual polymorphism through the occurrence of two (distyly) or three (tristyly) morph types that differ in floral traits (style length and anther position). We examined whether the long-styled (pin) and short-styled (thrum) morph types show differences in reproductive components and responses to habitat fragmentation in the distylous, self-incompatible perennial herb Primula veris. We documented reproductive components for pin and thrum individuals and their relationships with population size, plant density and morph ratio (pin frequency), in nine populations from Flanders (northern Belgium) located in fragmented habitats of the intensively used agricultural landscape. Seed abortion increased in small populations as a result of inbreeding depression. Fruit set increased with plant density. Seed set was positively related to pin proportion. Seed set was higher for pin than thrum in small populations, but lower in large populations. Two hypotheses can be considered to explain these morph-specific differences: a pollen transfer asymmetry, and a reproductive advantage for the partially self-compatible pin morph. Morph types appear to respond differently to habitat fragmentation constraints. A floral morph type showing partial self-compatibility may be favored in populations under pollination failure, because it can increase reproductive success and mating opportunities through intramorph crosses.

Quantifying deficits in the perception of fear and anger in morphed facial expressions after bilateral amygdala damage

Amygdala damage has been associated with impairments in perceiving facial expressions of fear. However, deficits in perceiving other emotions, such as anger, and deficits in perceiving emotion blends have not been definitively established. One possibility is that methods used to index expression perception are susceptible to heuristic use, which may obscure impairments. To examine this, we adapted a task used to examine categorical perception of morphed facial expressions [Etcoff, N. L., & Magee, J. J. (1992). Categorical perception of facial expressions. Cognition, 44(3), 227–240]. In one version of the task, expressions were categorized with unlimited time constraints. In the other, expressions were presented with limited exposure durations to tap more automatic aspects of processing. Three morph progressions were employed: neutral to anger, neutral to fear, and fear to anger. Both tasks were administered to a participant with bilateral amygdala damage (S.P.), age- and education-matched controls, and young controls. The second task was also administered to unilateral temporal lobectomy patients. In the first version, S.P. showed impairments relative to normal controls on the neutral-to-anger and fear-to-anger morphs, but not on the neutral-to-fear morph. However, reaction times suggested that speed-accuracy tradeoffs could account for results. In the second version, S.P. showed impairments on all morph types relative to all other subject groups. A third experiment showed that this deficit did not extend to the perception of morphed identities. These results imply that when heuristics use is discouraged on tasks utilizing subtle emotion transitions, deficits in the perception of anger and anger/fear blends, as well as fear, are evident with bilateral amygdala damage.

Time-domain signal divergence and discrimination without receptor modification in sympatric morphs of electric fishes

Polymorphism in an animal communication channel provides a framework for studying proximate rules of signal design as well as ultimate mechanisms of signal diversification. Reproductively isolated mormyrid fishes from Gabon's Brienomyrus species flock emit distinctive electric organ discharges (EODs) thought to function in species and sex recognition. Species boundaries and EODs appear congruent in these fishes, with the notable exception of three morphs designated types I, II and III. Within the species flock, these morphs compose a monophyletic group that has recently been called the magnostipes complex. Co-occurring morphs of this complex express distinctive EODs, yet they appear genetically indistinguishable at several nuclear loci. In this study, we investigated EOD discrimination by these morphs using both behavioral and physiological experiments. During the breeding season, wild-caught type I and type II males showed evidence that they can discriminate their own morph's EOD waveform from that of a sympatric and genetically distinct reference species. However, we found that type I and type II males exhibited an asymmetry in unconditioned responses to paired playback of EODs recorded from type I versus type II females. Males of the type II morph responded preferentially to EODs of type II females, whereas type I males did not appear to discriminate homotypic and heterotypic EODs in our experimental paradigm. Part of this behavioral asymmetry may have resulted from a previously undetected difference in adult size, which may have enhanced apparent discrimination by the smaller morph (type II) due to a relatively higher risk of injury from the larger morph (type I). Knollenorgan receptors, which mediate electrical communication in mormyrids, showed similar spectral tuning in type I and type II. These electroreceptors coded temporal features of any single magnostipes-complex EOD with similar patterns of time-locked spikes in both morphs. By contrast, Knollenorgans exhibited distinctive responses to different EOD waveforms. These results suggest that discrete EOD variation in this rapidly diversifying complex is functional in terms of morph-specific advertisement and recognition. Time-domain signal divergence has outpaced frequency-domain divergence between sympatric morphs, requiring little to no change in receptor response properties. We discuss our findings in light of a model for EOD time-coding by the Knollenorgan pathway, as well as evolutionary hypotheses concerning sympatric signal diversification in the magnostipes complex.

Dimorphic male midshipman fish: reduced sexual selection or sexual selection for reduced characters?

In most taxa with male dimorphisms, some males are large in body size with exaggerated secondary (exaggerated morph), whereas other males in the same population are small and have reduced secondary (reduced morph). What selective pressures cause male dimorphisms? Reduced morphologies may result when a) some males develop a morphology that, in the absence of selection pressures for an exaggerated morphology, reduces energetic and developmental costs and/or b) some males opt for an alternative morphology that does well at an alternative behavioral tactic such as cuckoldry. The 2 mechanisms could act together, but each alone is theoretically sufficient to drive dimorphisms. Here, we tested hypothesis b (sexual selection for reduced characters) in the plainfin midshipman fish, Porichthys notatus. Behavioral plasticity between territoriality and cuckoldry in an exaggerated male morph (type I) allows for a direct comparison of cuckoldry by exaggerated morph males to cuckoldry by reduced morph (type II) males. Compared with type I cuckolders, type II cuckolders were able to remain near the nest for longer periods before being chased by the territorial type I male, suggesting that the reduced type II morphology allows type II males to prolong the time before attack by territorial males. Combined with other studies showing a role of selection in maintaining the exaggerated morph, the data support the sexual selection for reduced characters hypothesis and elucidate how selection can act in different ways on different males to maintain 2 male morphologies within a single species. Copyright 2006.

Sequential ordering of morphed faces and facial expressions following temporal lobe damage

A card ordering task was developed to evaluate the role of the temporal lobe in perceiving subtle featural displacements of faces that contribute to judgments of facial expression and identity. Individuals with varying degrees of temporal lobe damage and healthy controls were required to manually sort cards depicting morphs of facial expressions or facial identities so that the cards were sequentially ordered from one morph endpoint to another. Four morph progressions were used—three emotion morphs (neutral-to-anger, neutral-to-fear, and fear-to-anger) and an identity morph. Five exemplars were given per morph type. Debriefing verified that participants were using feature-level cues to sort the cards. A patient with bilateral amygdala damage due to epilepsy did not differ in her sorting abilities from unilateral temporal lobectomy patients or controls. In contrast, a post-encephalitic patient with widespread left temporal lobe damage showed impairments that were most marked on the fear-to-anger and identity sorts. These results show that amygdala-damaged individuals can use information contained in facial expressions to solve tasks that rely on feature-level analysis, which recruits processing in other temporal lobe regions involved in making fine featural distinctions.

Within-population genetic variation in the distylous Primula veris: Does floral morph anisoplethy matter in fragmented habitats?

We studied genetic variation and structure in 24 populations of Primula veris , a distylous self-incompatible perennial herb that in Flanders (northern Belgium) occurs in fragmented habitats of the intensively used agricultural landscape. Distyly, which favours cross-fertilization, is characterized in this species by two genetically determined floral morph types (‘pin’ or ‘thrum’). Using 30 allozyme loci, we investigated within-population and within-morph genetic variation and structure in relation to distyly (stochastic morph bias and pin frequency), demography (population size, plant density), and evidence of recent bottlenecks. No significant relationship was found between genetic patterns and stochastic morph bias. However, F IS values were lower when pin frequency increases in the population. This may be explained by the partial self-compatibility of the pin morph, which, in conditions of pollination disruption, as observed for P. veris in fragmented habitats, may favour intra-pin outcrosses and increase mating opportunities in the population. A lower within-population structure can be expected because of a higher density of compatible mates and more random matings. Pin and thrum individuals showed similar levels of genetic variation. Three populations revealed a significant genetic differentiation between pin and thrum morphs ( F ST ⩾ 0.0 4 9 ), as a result of recent bottlenecks, spatial segregation of the morphs and/or disturbance of demographic processes. From our findings, we may hypothesize future changes in population mating processes in fragmented habitats, which might lead to pin excesses. However, due to slowing down life-history traits (long-lived perenniality and clonality), and demographic stochasticity randomly changing morph frequency, this may take time before being expressed. Evaluating the impact of morph frequencies on population dynamics is therefore essential for implementing conservation strategies of heterostylous plant species.

Floral and pollinator behaviour of flexistylous Bornean ginger, Alpinia nieuwenhuizii (Zingiberaceae)

Flexistyly is a unique floral mechanism involving extreme curving of the style. It was first described in Chinese ginger (Amomum, Zingiberaceae). This is a pioneer report on flexistylous gingers of Malesia, where most species of this family grow. We observed the floral behaviour and flower visitors in Alpinia nieuwenhuizii Val., a Bornean endemic. Although the floral behaviour and effective pollinators (carpenter bees, Xylocopa) were similar between the Bornean species and the previously reported flexistylous Alpinia, the pollinator behaviour between them strikingly differed with regard to the visit frequency of the pollinators showing a bimodal pattern during the day. This was a better match for the floral behaviour of the flexistylous Alpinia. Some gender differentiation observed between the two types of morphs is also discussed in the present study.

Differential effects of 11-ketotestosterone on dimorphic traits in a teleost with alternative male reproductive morphs

Species with dimorphic males typically have one morph (type I) associated with territoriality and courtship, and another morph (type II) associated with cuckoldry. In teleost fishes, type I males generally have higher levels of the androgen 11-ketotestosterone (KT) than type II males. Is KT causal to phenotypic differences between morphs? We investigated this question in the midshipman fish ( Porichthys notatus) in which type I males have detectable levels of KT during the breeding season, whereas levels in type II males are usually undetectable. Type I midshipman will either cuckold or hold territories and court females, whereas type II males are only known to cuckold. Acoustic courtship by type I's is supported by the sonic motor nucleus, which innervates a sound-producing sonic muscle. Type I males have larger sonic motor nuclei and larger sonic muscles than type II males, consistent with the more dynamic vocal repertoire of type I's. Here, we tested whether intraperitoneal KT implants in adult type II males would induce type I male-like traits in brain, sonic muscle, and behavior. Type II's treated with KT did not differ from blank-implanted type II's in sonic motor nucleus volume. Sonic muscle mass increased in KT-implanted type II's, but did not reach the relative mass naturally observed in type I's. While neither territoriality nor courtship were induced, cuckoldry behavior intensified in KT-implanted type II's. Thus, for some but not all characters, KT exaggerated the expression of already existing type II male traits rather than inducing the type I male-like traits of territoriality, courtship, and an expansive vocal motor system.

Mistakes in sexual recognition among sympatric Zygoptera vary with time of day and color morphism (Odonata: Coenagrionidae)

Abstract In odonates, female specific color polymorphisms appear to be an evolutionary response to sexual harassment, but we know little about the decision rules males use when searching for variable females. For two sympatric species of Enallagma, we measured male responses to live female variants under field conditions, early and later in the day. In the morning, when the operational sex ratio was the most male-biased and female density the lowest, males of the polymorphic E. civile did not discriminate among conspecific female morphs, and reacted sexually to the andromorphic females of E. aspersum, a monomorphic species. ‘Then, male E. aspersum did not favor conspecific females over E. civile morphs. Both morph types were more confusing for males than were conspecific male signals. However, after 13:00 h, males of both species made few mistakes, and E. civile males reacted sexually relatively less often to conspecific andromorphs, the minority morph in this population. The changes in a male's sexual re...

Labile male morphology and intraspecific male polymorphism in the Philotrypesis fig wasps

We investigate the evolution of male morphology in the fig wasps belonging to the genus Philotrypesis (Chalcidoidea, Sycorectinae). We first reconstruct the phylogenetic relationships of Philotrypesis associated with African figs using nuclear and mitochondrial DNA. We then determine male morphotypes in the species included in our phylogeny and show that intraspecific polymorphism is common. Most species present two types of males and some species have up to three types. These morphotypes are believed to represent alternative mating tactics: some males show morphological adaptations to fighting, others are winged dispersers and others are small sneakers. Mapping out these variations onto our phylogeny reveals that the combination of morphs changes randomly along the branches of the tree. Both parsimony and likelihood approaches indicate that there has been at least one transition from dimorphism to trimorphism, several gains and losses of the small morph and two independent acquisitions of the winged morph. Using maximum likelihood analyses of character evolution, we estimate transition rates for each morph and show that the evolution of each type of morph are not correlated and that forward and backward transition rates are not significantly different. Our results altogether suggest that male morphology is evolutionary labile, it responds quickly to selection imposed by the mating environment. This study, also suggests that seemingly complex phenotypes, such as winged males, can evolve several times and can even be recreated after having been lost.

Allozyme polymorphism and conservation of the Lake Kasumigaura population of Nymphoides peltata

A population of the perennial floating-leaved plant, Nymphoides peltata, in Lake Kasumigaura has declined rapidly in recent years, with no recruitment from seedlings. To obtain genetic information to facilitate a restoration program, we investigated allozyme polymorphisms in subpopulations from various parts of the lake. Only 14 multilocus genotypes were detected, although a relatively high percentage of polymorphic loci and high heterozygosity was present throughout the lake. Allozyme data indicated that the Lake Kasumigaura population consists of a single subpopulation that includes several clones consisting of two types of heterostylous floral morphs, and some subpopulations each consisting of a single clone. Preservation of the multiclonal subpopulation is essential for conserving the seed production and clonal diversity of N. peltata in Lake Kasumigaura.

Reduced reproductive success in small populations of the self‐incompatiblePrimula vulgaris

Summary 1 Habitat fragmentation and the resulting decline in population size can affect biotic interactions and reproductive success of plant species. We investigated the impact of habitat type, population size, morph type and frequency, plant density, floral display and predation on different reproductive components in 16 populations of the distylous self-incompatible perennial herb, Primula vulgaris , a rare, declining species in Belgium. 2 Although habitat type accounted for significant variation in population size, we did not find any relation between habitat type and either reproductive and vegetative characteristics. Population size, however, strongly affected reproductive success, such that plants in small populations produced significantly fewer fruits per plant and seeds per fruit, and therefore fewer seeds per plant. 3 No significant difference was found between morph types for any reproductive characteristic, nor an interaction with population size. However, when morph frequency was strongly biased ( ≥ 1 : 3), the proportion of flowers setting fruit and the number of seeds per fruit were significantly lower in individuals of the common morph type. 4 Within populations, individual plants varied tremendously in size and floral display. Total number of fruits per plant significantly increased with floral display, but the highest fruit set per flower was found at intermediate flower number. 5 The proportion of fruit suffering pre-dispersal predation per plant significantly increased with floral display, but this did not offset the potential fitness gains of producing a large display. Furthermore, the absolute number of predated fruits per plant was significantly and positively affected by the interaction of the total number of fruits per plant and the density of the population.

Geographic Variation in Shell Morphology of the Rock Shell, Thais clavigera (Gastropoda: Muricidae) According to Environmental Difference in Korean Coasts

Geographic variation in shell morphology of Thais clavigera <TEX>$(K\"{u}ster)$</TEX> (Gastropoda: Muricidae) was investigated using samples collected from 24 sites along the Korean coast. Multivariate statistical analysis was applied to 9 morphometric and 4 categorical variables. The shells of T. clavigera were classified into two distinct morph types (Type-W and -E). Temperature and salinity of the sampling sites were significantly correlated with the incidence of morph types. Relative abundance of Type-W (thin, yellowish brown shell with triangular nodules) was positively correlated with temperature and negatively correlated with salinity. In contrast, relative abundance of Type-E (thick, dark purple shell with round nodules) was negatively correlated with temperature and positively correlated with salinity. Possible correlation between environmental factors (temperature and salinity) and morphological variations in the shells were discussed.

Mate choice in divergent morphs of the gastropod mollusc Littorina saxatilis (Olivi): speciation in action?

We investigated mate choice in the gastropod Littorina saxatilis (Olivi) in the laboratory, using snails taken from two shores, 20km apart. The snails occur in two distinct shell morphs in the high (H) and mid (M) shore zones, known to show signs of a postzygotic reproductive barrier between them. We found that mating was assortative not only between the forms H and M on a single shore, but also between snails from the two different shores. Mating preference appeared to be based on morph type over this distance. The snails may be detecting and responding to an extrinsic attribute reflecting a common microhabitat (but across a distance of 20km), and/or an intrinsic attribute to do with the organisms themselves. These findings are further evidence of behaviour associated with the evolution of reproductive isolation in this gastropod species.

Differential ovule development following self‐ and cross‐pollination: the basis of self‐sterility in Narcissus triandrus (Amaryllidaceae)

Self-pollination results in significantly lower seed set than cross-pollination in tristylous Narcissus triandrus. We investigated structural and functional aspects of pollen-pistil interactions and ovule-seed development following cross- and self-pollination to assess the timing and mechanism of self-sterility. Ovule development within an ovary was asynchronous at anthesis. There were no significant differences in pollen tube behavior following cross- vs. self-pollination during the first 6 d of growth, regardless of style morph type. Double fertilization was significantly higher following cross- vs. self-pollination. Aborted embryo development was not detected following either pollination type up to seed maturity. Prior to pollen tube entry, a significantly greater number of ovules ceased to develop following self- vs. cross-pollination. These results indicate that self-sterility in N. triandrus operates prezygotically but does not involve differential pollen tube growth typical of many self-incompatibility (SI) systems. Instead, low seed set following self-pollination is caused by a reduction in ovule availability resulting from embryo sac degeneration. We hypothesize that this is due to the absence of a required stimulus for normal ovule development. If this is correct, current concepts of SI may need to be broadened to include a wider range of pollen-pistil interactions.

EVOLUTION OF A POLYMORPHISM FOR OUTCROSSING RATE IN SENECIO VULGARIS: INFLUENCE OF GERMINATION BEHAVIOR.

A difference in germination behavior between the highly selling, nonradiate variant of Senecio vulgaris and the more outcrossing, radiate form had a large effect on the relative female fitness of these two morphs raised in mixed stands under conditions similar to those experienced in the wild. Of particular significance was the finding that female fitness differences between morphs were reversed in early- and late-sown plots. This was because a greater proportion of nonradiate than radiate seed germinated directly after sowing, and in early-sown plots a relatively large proportion of early-germinated nonradiate seedlings survived winter to produce large, highly fecund plants the following summer, that contributed greatly to the total female fitness of the nonradiate morph. In contrast, in late-sown plots (i.e., sown two weeks later), survivorship of early germinated seedlings was much reduced, and the radiate morph had an advantage because most radiate seed delayed germination until spring, therefore avoiding seedling mortality during winter. The effect of the association between morph type and germination behavior on morph female fitness in S. vulgaris is clearly important in regard to the evolution of the polymorphism for outcrossing rate in the species. This, in turn, emphasizes the point that an understanding of factors responsible for the evolution and maintenance of polymorphisms for outcrossing rate in the wild should be based on a detailed examination of the ecological genetics of such polymorphisms that extend beyond traditional studies of pollen discounting and inbreeding depression.

SELECTION ON THE COLOR POLYMORPHISM IN HAWAIIAN HAPPY-FACE SPIDERS: EVIDENCE FROM GENETIC STRUCTURE AND TEMPORAL FLUCTUATIONS.

Throughout this century genetic polymorphisms for color have been widely used as a research tool to allow insights into key evolutionary processes. Although color variants can often be diverse within populations, frequencies of different morphs may be similar across populations, either as a result of balancing selection or gene flow. Under these circumstances selection can be extremely difficult to demonstrate. Here we test for balancing selection on the naturally occurring color forms of the Hawaiian happy-face spider, Theridion grallator with two approaches. First, allozyme loci are used to generate a null model against which to test selection. Frequencies of alleles involved in the color polymorphism of T. grallator are used to generate another estimate for comparison. The results suggest that statistically similar frequencies of color morphs among populations of T. grallator may be maintained by some form of balancing selection. Second, we make use of an unusual event in which the normally stable frequencies of unpatterned and patterned morphs within a population were found to have shifted toward an excess of unpatterned morphs. We scored offspring of all fertilized, unpatterned (bottom-recessive) females found during this period of skewed morph frequencies and also in a year when morph frequencies were normal to deduce paternal color phenotypes. Mating was found to be random in the normal year, but in the perturbed year females had mated with rare (patterned) males twice as frequently as expected on the basis of the frequency of this morph type in the population. Both of these results are consistent with selection operating on the color polymorphism, and we speculate that apostatic selection, perhaps mediated by bird predators, may provide the mechanism.

Dimorphic Males Display Alternative Reproductive Strategies in the Marine Amphipod Jassa marmorata Holmes (Corophioidea: Ischyroceridae)

AbstractDiscrete dimorphism of males within a species is often the result of selection for alternative reproductive strategies, and these strategies may be evolutionarily stable (ESS). An ESS may be either mixed (genetically fixed differences) or conditional (flexible differences related to varying environmental conditions) (PARKER 1984). Mature males of the marine amphipod Jassa marmorata are dimorphic. Large ‘major’ males have a greatly enlarged thumb (propodus) on their 2nd gnathopods, while small ‘minor’ males exhibit thumbs that are reduced, and CONIAN (1989) suggested that minors may exhibit a different mating strategy from majors. Ratios of males and females fluctuate seasonally (FRANZ 1989) and female body size is inversely correlated with temperature (FRANZ 1989) so male dimorphism could be a flexible response to varying environmental conditions. I sampled a natural population of J. marmorata over a 1‐yr period, quantified major and minor morphology, and measured male behaviour and mating success in experimental arenas that contained varying proportions of male morphs and females. Morphology of the two morphs is discrete; female body size varies with season with significantly smaller individuals in summer and fall; body size predicts morph type; and ratios of majors, minors and females fluctuate seasonally. Finally I showed that majors and minors use different mating tactics to gain access to receptive females, and that these behaviours depend on the male's own morphology and on the environmental setting that it finds itself in. Major males fight, display and attempt to evict other males to mate with receptive females. Minors never fight with, display to or attempt to evict majors, but they infrequently display to and attempt to evict other minor males. Furthermore, mating success of the two morphs was not significantly different and may depend on whether males are with a majority or minority of their own type. These data support the conclusion that major and minor male J. marmorata display evolutionarily stable alternative reproductive strategics, but more work should address the nature of this ESS.

Polymorphism in the Ostertagiinae Lopez-Neyra, 1947 and comments on the systematics of these nematodes

A list of 19 polymorphic species of the generaOstertagia, Orloffia, Teladorsagia, Marshallagia andSpiculopteragia is presented. The morphs of the polymorphic species in question have identical females and show similarities in some male features, e.g. the structure of the oesophagus, the synlophe and the shape of the rays of the copulatory bursa. However, they differ by characters which are recognised as generic features, e.g. the structure of spicules, gulbernaculum and genital cone. The presence or lack of minor morphs is a generic feature in the Ostertagiinae. Each polymorphic genus has a defined type of minor morph.