Poeciliid Fish Research Articles

Allelic expression of IGF2 in live-bearing, matrotrophic fishes

The parental conflict, or kinship, theory of genomic imprinting predicts that parent-specific gene expression may evolve in species in which parental investment in developing offspring is unequal. This theory explains many aspects of parent-of-origin transcriptional silencing of embryonic growth regulatory genes in mammals, but it has not been tested in any other live-bearing, placental animals. A major embryonic growth promoting gene with conserved function in all vertebrates is insulin-like growth factor 2 (IGF2). This gene is imprinted in both eutherians and marsupials, as are several genes that modulate IGF2 activity. We have tested for parent-of-origin influences on developmental expression of IGF2 in two poeciliid fish species, Heterandria formosa and Poeciliopsis prolifica, that have evolved placentation independently. We found IGF2 to be expressed bi-allelically throughout embryonic development in both species.

Variation between and within the sexes in body size preferences

Although it is often assumed that males and females have mating preferences for larger individuals of the other sex, potential underlying differences between male and female preferences for body size are not commonly investigated. Here, sexual differences in body size preferences are examined in the poeciliid fish, Brachyrhaphis rhabdophora. Females preferred larger males to smaller males, but preference did not appear to be affected by female size. One population-level analysis for males did not indicate an overall preference for larger females. A closer examination, however, revealed an effect of male size on preference; larger males preferred larger females, while smaller males preferred smaller females. It appears then that females, regardless of size, share a preference for large males, but males differ in their behaviour, depending on their body size. In addition, while the degree of difference in size between paired females did not appear to affect male preference, the degree of difference in size between paired males strongly affected female preference; the greater the difference, the more strongly females preferred the larger male. Thus, intersexual selection is found to operate in both sexes, but how it operates appears to differ. Intrasexual and intersexual differences in mating behaviour may be missed when evaluating population-wide preferences. That is, there can be underlying differences in how the sexes respond and the consequences of such differences should be considered when investigating mate choice. The results are considered in terms of the evolution of mating preferences, alternative mating strategies, assortative mating, the maintenance of trait variation in a population, and current methods to evaluate mating preferences.

Variation between and within the sexes in body size preferences

Although it is often assumed that males and females have mating preferences for larger individuals of the other sex, potential underlying differences between male and female preferences for body size are not commonly investigated. Here, sexual differences in body size preferences are examined in the poeciliid fish, Brachyrhaphis rhabdophora . Females preferred larger males to smaller males, but preference did not appear to be affected by female size. One population-level analysis for males did not indicate an overall preference for larger females. A closer examination, however, revealed an effect of male size on preference; larger males preferred larger females, while smaller males preferred smaller females. It appears then that females, regardless of size, share a preference for large males, but males differ in their behaviour, depending on their body size. In addition, while the degree of difference in size between paired females did not appear to affect male preference, the degree of difference in size between paired males strongly affected female preference; the greater the difference, the more strongly females preferred the larger male. Thus, intersexual selection is found to operate in both sexes, but how it operates appears to differ. Intrasexual and intersexual differences in mating behaviour may be missed when evaluating population-wide preferences. That is, there can be underlying differences in how the sexes respond and the consequences of such differences should be considered when investigating mate choice. The results are considered in terms of the evolution of mating preferences, alternative mating strategies, assortative mating, the maintenance of trait variation in a population, and current methods to evaluate mating preferences.

Multiple paternity and population genetic structure in natural populations of the poeciliid fish, Heterandria formosa.

Heterandria formosa Agassiz, exhibits internal fertilization, internal brooding of embryos, sperm storage and an extreme level of superfetation. In this study we used microsatellite markers to examine variation among seven populations that exhibited significant variance in their histories of population density. We found that the populations were genetically distinct and that the heterozygosity increased as population density increased. We also examined paternity in three of those populations and found that the number of sires per female and the number of sires per brood increased with population density. Overall, the rates of multiple paternity are quite low relative to other species. The correlations with population density suggest that contact rates play a critical role in the breeding system in this species but the low rates of multiple paternity suggest that females may exert control over fertilization of their ova.

Seasonal and yearly population dynamics of two exotic helminths, Camallanus coti (Nematoda) and Bothriocephalus acheilognathi (Cestoda), parasitizing exotic fishes in Waianu Stream, O'ahu, Hawaii.

Exotic poeciliid fishes introduced into Hawaiian freshwaters are responsible for the introduction of several exotic parasites, of which the most important are Camallanus cotti and Bothriocephalus acheilognathi in terms of potential disease threat to native stream fishes. This roundworm and tapeworm are the most prevalent and abundant freshwater fish helminths in Hawaiian streams. This study examined the seasonal and yearly population structure of C. cotti and B. acheilognathi to determine if the tropical Hawaiian environment characterized by low climatic variability permits continuous opportunities for parasite transmission regardless of time of year. Camallanus cotti displayed seasonal differences in prevalence and mean abundance, whereas B. acheilognathi did not. Camallanus cotti prevalence and mean abundance were higher in the Hawaiian summer (47.7%, 0.79) than in winter (25.8%, 0.36). A seasonal relationship of C. cotti levels is likely explained by extensive rains associated with the Hawaiian winter season, which may act to decrease parasite transmission by flushing infected poeciliid hosts, intermediate copepod hosts, and possibly free-living infective worm stages downstream. Bothriocephalus acheilognathi displayed low prevalence and mean abundance in both summer (4.0%, 0.06) and winter (6.5%, 0.07), and it may be difficult to detect seasonal changes due to these low levels. Camallanus cotti prevalence and mean abundance remained relatively constant from the summer of 1995 to the summer of 1999, indicating that levels of this roundworm are stable in Waianu Stream. Whereas B. acheilognathi prevalence and mean abundance were low during the summer of 1995 and the summer of 1997, a dramatic peak in prevalence and mean abundance was observed in the summer of 1998 (41.2%, 1.06), with levels decreasing sharply in the summer of 1999 (4.4%, 0.07). It appears that B. acheilognathi also is present in stable populations at low levels, even though levels rose sharply during a single year.

Critique of parsimony analysis of endemicity as a method of historical biogeography

AbstractAim Assess the value of parsimony analysis of endemism as either an a priori (cladistic) and an a posteriori (phylogenetic) method of historical biogeography.Location World‐wide.Methods Parsimony analysis of endemicity (PAE) and Brooks parsimony analysis (BPA).Results Parsimony analysis of endemicity is capable of finding correct and unambiguous area relationships only under scenarios of vicariance in combination with non‐response to vicariance or extinction. An empirical comparison between PAE and BPA, using the poeciliid fish genera Heterandria and Xiphophorus, demonstrates that PAE fails to document much of the historical complexity in this relatively simple system.Main conclusions The a priori assumptions of PAE are far more restrictive than those made by other a priori methods, limiting its utility as a method of cladistic biogeography. The inability of PAE to detect perfect vicariance or biogeographical histories involving dispersal, renders it unsuitable as a method of phylogenetic biogeography.

Disturbed sexual characteristics in male mosquitofish (Gambusia holbrooki) from a lake contaminated with endocrine disruptors.

Previous laboratory studies have demonstrated that estrogenic and antiandrogenic chemicals can alter several sexual characteristics in male poeciliid fishes. Whether similar disturbances occur under field conditions remains to be confirmed. Lake Apopka, Florida, is contaminated with numerous chemicals, some of which possess endocrine-disrupting activity. Male mosquitofish (Gambusia holbrooki) were collected monthly from December 2000 through May 2001 from Lake Apopka and two nearby reference lakes, Orange Lake and Lake Woodruff National Wildlife Refuge. Selected sexual characteristics were compared temporally and among lakes during the collection period. Male fish from Lake Apopka had slightly shorter gonopodia and on average 32 and 47% fewer sperm cells per milligram testis, when compared with the fish collected from Orange Lake and Lake Woodruff, respectively. The testes weights increased markedly during spring, with significantly smaller testes in fish from Lake Apopka than from Orange Lake, but surprisingly, the smallest testes occurred in males obtained from the Lake Woodruff population. The highest liver weights were found in the Lake Apopka population. Whole-body concentrations of testosterone and estradiol varied among months; the peak testosterone concentration occurred in January and was significantly lower in male fish from Lake Apopka compared with Orange Lake. The intensity of male courtship behavior was highly correlated to body testosterone concentration, but no statistically significant differences in sexual behavior among the lakes were found. We conclude that sexual characteristics of relevance to male reproductive capacity are altered in the Lake Apopka mosquitofish population, and we discuss the presence of chemicals with antiandrogenic effects in Lake Apopka as a possible cause of the observed alterations.

Sex Chromosomes and Sexual Selection in Poeciliid Fishes

We propose that the evolution of female preferences can be strongly influenced by linkage of attractive male traits to the Y chromosome and female preferences to the X chromosome in male heterogametic species. Such linkage patterns are predicted by models of the evolution of sexually antagonistic genes. Subsequent recombination of attractive male characters from the Y to the X would create physical linkage between attractive male trait and preference. A literature survey shows that Y linkage of potentially sexually antagonistic traits is common in poeciliid fishes and other species with sex chromosomes that are not well differentiated, but may also occur in taxa with degenerate Y chromosomes. In the guppy, attractive male traits are primarily Y and X linked; a literature review of the inheritance of sex-limited attractive male characters suggests that 16 are Y linked, 24 recombine between the X and Y, two are X linked, and two are autosomal. Crosses and backcrosses between high female preference (Endler's live-bearers) and low female preference (Rio San Miguel) guppy populations show that this character has a strong additive genetic component and that it will be possible to investigate the physical linkage of male and female sexually selected characters in this species through mapping studies.

Congruence between the sexes in preexisting receiver responses

Preexisting receiver biases have been shown to affect how females detect and respond to new conspecific traits in a mate choice context. Although preexisting biases have often been discussed in the context of female mate choice, these biases need not be sex limited. In the genus Xiphophorus, swordtail males possess a sexually selected trait, the sword. Here I consider evidence that the state of a bias favoring sworded conspecifics may be generally shared by the sexes in taxa in which the sword has not arisen. In three unsworded species of poeciliid fishes, both males and females prefer members of the other sex with swords. In a fourth species, males and females share the absence of a response to a sword. This congruity between the sexes suggests that response biases may not be sex limited and that the sexes could historically share common mechanisms producing shared mating responses. Alternatively, selection may tend to result in parallel changes in biases in the sexes. This work expands our understanding of receiver biases by using a phylogenetic approach to examine whether biases are historically shared by the sexes and suggests that there can be general congruence between the sexes in such biases.

Geographical variation in male genitalia inBrachyrhaphis episcopi(Poeciliidae): is it sexually or naturally selected?

Male poeciliid fishes inseminate females using an intromittent organ called the gonopodium. Here we report on natural variation in gonopodium size both within and between 12 populations of the freshwater fishBrachyrhaphis episcopi(Poeciliidae) in Panama. We show that males from sites with more predatory fish species have, on average, a relatively longer gonopodium than males inhabiting sites with fewer predatory fish. Gonopodium length was not correlated with the site‐specific adult sex ratio and the average sex ratio was more strongly female biased at sites with more predatory fish. The gonopodium exhibited lower phenotypic variance than the average for sexually selected traits and it generally showed negative allometry. Our results are similar to those reported for the guppyPoecilia reticulata. Two alternative hypotheses for these findings are discussed. First, that population differences are sexually selected. Second, that they are an incidental consequence of environmental differences between sites. Specifically, that higher water flow rates select for enlarged fin size and stockier bodies in downstream sites where predatory fish are more common.

Female discrimination against sworded males in a poeciliid fish

Female poeciliid fish of the sister genera Xiphophorus and Priapella share a mating preference favouring a sword, despite phylogenetic evidence that the sword arose in Xiphophorus. This conclusion is based on the presence of a preference favouring a sword in three unsworded species and one sworded species. Here I investigate the response to a sword in the poeciliid fish Heterandria bimaculata by testing females with conspecific males that differ in the presence or absence of a sword. Female H.bimaculata were found to prefer unsworded males, thus sworded males were discriminated against. I also tested the strength of female response; females did not show a differential response to males based on sword length. The preference for unsworded males could indicate that when new male traits arise, biases might oppose trait evolution, even in cases in which similar male traits in related taxa are favoured. Explanations for the female preference for unsworded males are discussed.

Repeatable female preferences, mating order and mating success in the poeciliid fish, Heterandria formosa

Sexual selection via female choice can afford preferred males comparably higher mating success than those males that lack preferred traits. In addition, many models of sexual selection assume that both male traits and female preferences are heritable. In this study we test whether females of the poeciliid fish, Heterandria formosa, have repeatable pre-copulatory preferences for larger males. We also test whether female pre-copulatory preferences are always reliable indicators of male mating success. When given a choice between a large and a small male, females prefer larger males, and the repeatability of this preference is high. Although there are no overall differences in male mating success between large and small males, large males have a higher mating success when they are the first to mate than when they are the second to mate. Likewise, preferred males also have higher mating success when they are the first to mate than when they mate second. Therefore, the repeatable female preferences observed in this study only predict male mating success when the preferred male mates first. These results illustrate that even significantly repeatable female preferences do not translate into male mating success, which is an assumption of many examinations of the importance of female choice in sexual selection.

Assessing Modes of Speciation: Range Asymmetry and Biogeographical Congruence

Lynch [1989, in“Speciation and Its Consequences” (D. Otte and J. A. Endler, Eds.), pp. 527–553, Sinauer, Sunderland, MA, proposed a methodology for assessing the frequency of occurrence of various modes of initiating species formation, using a combination of phylogenetic relationships and relative size of geographic ranges for sister species and sister groups. Historical biogeography provides an alternative criterion for assessing modes of species formation. Species whose distributions conform to a general (replicated in multiple clades) pattern of area relationships are deemed to be the result of vicariance (including microvicariance) regardless of the details of current geographic range. Species exhibiting unique biogeographic distributions are the result of peripheral isolates speciation and postspeciation dispersal. Lineage duplications indicate sympatric speciation. An empirical assessment of this alternative approach was performed using the most recent phylogenetic trees and geographical distribution data on the Mesoamerican poeciliid fish comprising the genera Xiphophorus and Heterandria. The single area cladogram produced by secondary Brooks Parsimony Analysis indicates 3 vicariant events (accounting for seven extant species and the common ancestor of the northern swordtails, which are not further analyzed) and at least 13 episodes of peripheral isolates speciation. Two of the 10 areas considered in previous analyses are vicariant areas of endemism, 1 is historically unique due to a single episode of peripheral isolates speciation, and the remaining 7 have reticulated histories of speciation. The results corroborate inferences of speciation modes made by Lynch for the same data.

Assessing Modes of Speciation: Range Asymmetry and Biogeographical Congruence

Lynch [1989, in “Speciation and Its Consequences” (D. Otte and J. A. Endler, Eds.), pp. 527–553, Sinauer, Sunderland, MA, proposed a methodology for assessing the frequency of occurrence of various modes of initiating species formation, using a combination of phylogenetic relationships and relative size of geographic ranges for sister species and sister groups. Historical biogeography provides an alternative criterion for assessing modes of species formation. Species whose distributions conform to a general (replicated in multiple clades) pattern of area relationships are deemed to be the result of vicariance (including microvicariance) regardless of the details of current geographic range. Species exhibiting unique biogeographic distributions are the result of peripheral isolates speciation and postspeciation dispersal. Lineage duplications indicate sympatric speciation. An empirical assessment of this alternative approach was performed using the most recent phylogenetic trees and geographical distribution data on the Mesoamerican poeciliid fish comprising the genera Xiphophorus and Heterandria. The single area cladogram produced by secondary Brooks Parsimony Analysis indicates 3 vicariant events (accounting for seven extant species and the common ancestor of the northern swordtails, which are not further analyzed) and at least 13 episodes of peripheral isolates speciation. Two of the 10 areas considered in previous analyses are vicariant areas of endemism, 1 is historically unique due to a single episode of peripheral isolates speciation, and the remaining 7 have reticulated histories of speciation. The results corroborate inferences of speciation modes made by Lynch for the same data.

Sex Chromosomes and Sexual Selection in Poeciliid Fishes

We propose that the evolution of female preferences can be strongly influenced by linkage of attractive male traits to the Y chromosome and female preferences to the X chromosome in male heterogametic species. Such linkage patterns are predicted by models of the evolution of sexually antagonistic genes. Subsequent recombination of attractive male characters from the Y to the X would create physical linkage between attractive male trait and preference. A literature survey shows that Y linkage of potentially sexually antagonistic traits is common in poeciliid fishes and other species with sex chromosomes that are not well differentiated, but may also occur in taxa with degenerate Y chromosomes. In the guppy, attractive male traits are primarily Y and X linked; a literature review of the inheritance of sex‐limited attractive male characters suggests that 16 are Y linked, 24 recombine between the X and Y, two are X linked, and two are autosomal. Crosses and backcrosses between high female preference (Endler’s live‐bearers) and low female preference (Rio San Miguel) guppy populations show that this character has a strong additive genetic component and that it will be possible to investigate the physical linkage of male and female sexually selected characters in this species through mapping studies.

Phylogeny of Limia (Teleostei: Poeciliidae) Based on NADH Dehydrogenase Subunit 2 Sequences

Complete sequences for the mitochondrial gene NADH Dehydrogenase 2 (ND2) and partial sequences for the tRNA-Met and tRNA-Trp genes were obtained for 11 populations of the poeciliid fish genus (or subgenus) Limia, including species from Hispaniola, Cuba, Jamaica, and Grand Cayman Islands. Additional sequences for Limia (2 species), Pamphorichthys (3), Poecilia (16), and Xiphophorus (1), all from the tribe Poeciliini, were extracted from GenBank, as was a sequence for Heterandria formosa, from the tribe Heterandriini. Phylogenetic analyses included parsimony, distance methods, and maximum-likelihood. Several Limia species groups that had been proposed based on morphological data were evaluated. The versicolor species group was strongly supported, as was the close relationship between the Cuban Limia vittata and the Grand Cayman Limia caymanensis. However, the proposed vittata species group was not upheld by the ND2 data. The phylogenetic position of Limia melanogaster, from Jamaica, was either sister to the versicolor species group or basal to all other included Limia species. Limia was found to be monophyletic; however, Limia species from the island of Hispaniola were not monophyletic. There was little support for any proposed sister group to Limia. The phylogeny was used to reconsider a previous comparative study of poeciliid courtship behavior and sexual dimorphism. The data indicated that there may have been two independent appearances of courtship display behavior in Limia; considering the tribe Poeciliini as a whole, there is evidence for as many as five appearances of display (including two within Limia), or a complex pattern of gains and losses of such behavior. The application of phylogenetic information to the comparative study did not refute the previously hypothesized correlation between the presence of a courtship display and the presence of sexual dimorphism in poeciliid fish.

The Suitability of Calcein to Mark Poeciliid Fish and a New Method of Detection

The suitability of calcein as a marker for population studies depends on (1) the assumption that marked individuals have unaltered viability, (2) the fidelity of the calcein label, and (3) the facility with which calcein can be used. We examined the effects of calcein on survival, growth, and the timing and size at sexual maturity of least killifish Heterandria formosa and present a new method for detecting calcein. To test the assumption that marked individuals have unaltered viability, juvenile least killifish were immersed for 24 h in a 250-mg/L solution of calcein. A control group of same-aged juveniles was immersed in the same volume of water for 24 h without calcein. All individuals were then removed and reared individually in separate containers. Upon examination under an epifluorescent microscope, all calcein-treated fish showed fluorescent green marks on their scales and fin rays, whereas controls did not. The calcein treatment had no significant effect on growth and survival through 9 weeks nor on the age and size at maturity. We also designed a portable detector (uses InGaN blue LEDs as a light source) for distinguishing calcein-marked individuals; using either this new detector or a standard epifluorescent microscope, the fluorescent mark was visible on the test fish for up to 5 weeks in the laboratory, although some individuals required remarking (due to fading) at 2–3 weeks postimmersion. The calcein tag was also visible in the vertebrae of ethanol-preserved specimens for up to 6 years, provided specimens were stored in the dark.

Variability of genetic sex determination in poeciliid fishes.

Poeciliids are one of the best-studied groups of fishes with respect to sex determination. They present an amazing variety of mechanisms, which span from simple XX-XY or ZZ-ZW systems to polyfactorial sex determination. The gonosomes of poeciliids generally are homomorphic, but very early stages of sex chromosome differentiation have been occasionally detected in some species. In the platyfish Xiphophorus maculatus, gene loci involved in melanoma formation, in different pigmentation patterns and in sexual maturity are closely linked to the sex-determining locus in the subtelomeric region of the X- and Y- chromosomes. The majority of traits encoded by these loci are highly polymorphic. This phenomenon might be explained by the high level of genomic plasticity apparently affecting the sex-determining region, where frequent rearrangements such as duplications, deletions, amplifications, and transpositions frequently occur. We propose that the high plasticity of the sex-determining region might explain the variability of sex determination in Xiphophorus and other poeciliids.

Female mate choice in a mating system dominated by male sexual coercion

In poeciliid fishes, males can gain copulation either by courting females or through sexual coercion. In some species these two tactics coexist. However, in about half of the poeciliids, males do not display, females never cooperate during copulation and all matings are achieved by thrusting the intromittent organ toward the genital pore of apparently unaware females. In one of these species, the eastern mosquitofish (Gambusia holbrooki), the probability of insemination is influenced by the time females are previously deprived of males, suggesting that females exert some control over the occurrence of mating even in a system apparently dominated by sexual coercion. In the present study we investigated the tendency of female mosquitofish to approach males in relation to their reproductive status and the time they were previously deprived of males. The tendency to approach males increased in females that were previously deprived of males and in females that had recently given birth. When allowed to choose between males, male-deprived females preferred larger males and normally pigmented over melanistic males. Females preferred groups of three males over a single male, whereas the preference for three males over a group of one male and two females was only marginally significant. Collectively, these results suggest that, even when coercive mating is the only tactic adopted by males, females may be able to influence the outcome of these attempts, and thus exert some control over the paternity of their offspring. Key words: sexual selection, female choice, sexual coercion, Gambusia holbrooki, fish, mating tactics. [Behav Ecol 12:59–64 (2001)]

Evaluating the Effects of Introduced Rainbow Trout (Oncorhynchus Mykiss) on Native Stream Insects on Kauai Island, Hawaii; Contribution No. 2001-012 to the Hawaii Biological Survey, Bishop Museum

Rainbow trout (Oncorhynchus mykiss) and other salmonids have been widely stocked into upland streams throughout the world to provide a basis for sport fisheries, but the effects of such introductions on indigenous and endemic aquatic insect assemblages are poorly documented. In this study, we examine the impact of rainbow trout on the indigenous and endemic entomofauna of upland streams in Kokee State Park, Kauai, Hawaii, with particular emphasis on the potential threat trout pose to populations of endemic damselflies in the genus Megalagrion. Rainbow trout were introduced into the upland streams of Kauai beginning in the 1920s, with over 60 years of subsequent restocking. This study indicates, however, that streams in this area still maintain diverse populations of Megalagrion damselflies and other indigenous and endemic aquatic insects, both in catchments containing naturally reproducing trout populations and in catchments lacking rainbow trout. Our results indicate that the indigenous and endemic aquatic insect communities in the streams under study compare favorably in terms of density and taxonomic richness with other isolated and unimpacted streams elsewhere in Hawaii, and retain high densities and relative percentages of indigenous and endemic aquatic insect taxa. Our results demonstrate that the threats posed by conspicuous introduced species such as trout should not simply be assumed a priori on the basis of postulated negative interactions, because this may divert limited resources from programs aimed at control of other, potentially more destructive introduced taxa such as inconspicuous poeciliid fishes.