Poeciliid Fish Research Articles

Morphological and molecular description of eight new species of Gyrodactylus von Nordmann, 1832 (Platyhelminthes: Monogenea) from poeciliid fishes, collected in their natural distribution range in the Gulf of Mexico slope, Mexico

Eight new species of Gyrodactylus are described from Poecilia mexicana, Poeciliopsis gracilis, Pseudoxiphophorus bimaculatus [syn. = Heterandria bimaculata], and Xiphophorus hellerii collected in the Nautla and La Antigua River Basins in Veracruz, and in the Tecolutla River Basin in Puebla, Mexico. Analyzing the morphology of the marginal hooks, Gyrodactylus pseudobullatarudis n. sp. and Gyrodactylus xtachuna n. sp. are both very similar to Gyrodactylus bullatarudis; Gyrodactylus takoke n. sp. resembles Gyrodactylus xalapensis; Gyrodactylus lhkahuili n. sp. is similar to Gyrodactylus jarocho; and both Gyrodactylus microdactylus n. sp. and Gyrodactylus actzu n. sp. are similar to Gyrodactylus poeciliae in that all three species possess extremely short shaft points. A hypothesis of the systematic relationships of the eight new Gyrodactylus species and some of the known gyrodactylids infecting poeciliids was constructed with sequences of the Internal Transcribed Spacers (ITS1 and ITS2) and the 5.8S ribosomal gene of the rRNA. Phylogenetic trees showed that the new and previously described species of Gyrodactylus infecting poeciliid fishes do not form a monophyletic assemblage. Trees also showed that the eight new species described morphologically correspond to well-supported monophyletic groups; and that morphologically similar species are also phylogenetically close. Additionally, we correct previous erroneous records of the presence of Gyrodactylus bullatarudis on wild Poecilia mexicana and Xiphophorus hellerii collected in Mexico, as re-examination of the original specimens indicated that these corresponded to Gyrodactylus pseudobullatarudis n. sp. (infecting Poecilia mexicana and Xiphophorus hellerii) and to Gyrodactylus xtachuna n. sp. (on Xiphophorus hellerii). Finally, given the widespread anthropogenic translocation of poeciliid fishes for the aquarium trade and mosquito control programs, as well as the existence of invasive, feral poeciliid populations worldwide, we discuss the possibility that gyrodactylid parasites could be introduced along with the fish hosts-this work provides taxonomic information to assess that possibility, as it describes parasites collected from poeciliid fishes within their native distribution range.

The mate choice mind: studying mate preference, aversion and social cognition in the female poeciliid brain

Male phenotypes vary across mating systems, but what about the female mind? In poeciliid fishes, we observe that female response towards males varies, both at inter- and intraspecific levels. By using the natural variation in poeciliid male reproductive phenotypes, we can probe female responses in different social contexts to isolate the neuromolecular components associated with mate choice. Female Xiphophorus nigrensis exhibit differential nonapeptide (isotocin, vasotocin) gene expression between social contexts that differ by social affiliation; while differential expression of synaptic plasticity genes (neuroserpin, neuroligin-3, NMDAR) is more strongly predictive of social contexts that differ by mate preference discrimination. Gene expression and pharmacological manipulation with X. nigrensis females suggests that the social cognition task of mate choice involves neuromolecular processes linked to learning at the cellular (synaptic plasticity genes) and regional (amygdala and hippocampus) levels. Comparative assays between two poeciliid species with different mating systems provide the first hint of a shared neuromolecular pathway underlying female mate preference response. Yet, it is still unclear how specific components of this neuromolecular pathway (nonapeptides and synaptic plasticity processes) interact to coordinate behavioural tasks associated with mate choice encounters (vigilance, affiliation, evaluation and discrimination). Nevertheless, this collection of studies, coupled with the natural diversity of poeciliid mating systems, pave the way for exciting new insights into the social cognition of mate choice.

Transcriptomics of two evolutionary novelties: how to make a sperm-transfer organ out of an anal fin and a sexually selected "sword" out of a caudal fin.

Swords are exaggerated male ornaments of swordtail fishes that have been of great interest to evolutionary biologists ever since Darwin described them in the Descent of Man (1871). They are a novel sexually selected trait derived from modified ventral caudal fin rays and are only found in the genus Xiphophorus. Another phylogenetically more widespread and older male trait is the gonopodium, an intromittent organ found in all poeciliid fishes, that is derived from a modified anal fin. Despite many evolutionary and behavioral studies on both traits, little is known so far about the molecular mechanisms underlying their development. By investigating transcriptomic changes (utilizing a RNA-Seq approach) in response to testosterone treatment in the swordtail fish, Xiphophorus hellerii, we aimed to better understand the architecture of the gene regulatory networks underpinning the development of these two evolutionary novelties. Large numbers of genes with tissue-specific expression patterns were identified. Among the “sword genes” those involved in embryonic organ development, sexual character development and coloration were highly expressed, while in the gonopodium rather more morphogenesis-related genes were found. Interestingly, many genes and genetic pathways are shared between both developing novel traits derived from median fins: the sword and the gonopodium. Our analyses show that a larger set of gene networks was co-opted during the development and evolution of the “older” gonopodium than in the “younger,” and morphologically less complex trait, the sword. We provide a catalog of candidate genes for future efforts to dissect the development of those sexually selected exaggerated male traits in swordtails.

Coevolution of male courtship and sexual conflict characters in mosquitofish

Coerced copulations cause sexual conflict: they enhance male fitness at a potential cost to female fitness. In contrast, male display allows female mate choice and stimulates cooperation, and so is thought to be an alternative to coercion. Here, we investigate the evolutionary relationship between coercion and display in Gambusia, a genus of poeciliid fish. We used a novel Markov chain method to quantify the transition rate from a male display to a copulatory thrust (the display-to-thrust [DT] rate), which we use to measure the importance of male display to copulation. We assess the potential for male coercion and the female response to coercion based on morphology. We found positive relationships between DT rate and 2 morphologies that are indicative of coercion: male gonopodium shape and the male morphological mating advantage (the ratio of gonopodium shape to female papilla size scores). We suggest that male display can cause the evolution of traits involved in sexually antagonistic coevolution, or conversely that display can evolve as a byproduct of sexual antagonism.

The life cycle of Pseudosellacotyla lutzi (Digenea: Cryptogonimidae), in Aylacostoma chloroticum (Prosobranchia: Thiaridae), and Hoplias malabaricus (Characiformes: Erythrinidae), in Argentina.

Pseudosellacotyla lutzi ( Freitas, 1941), at present included in the Faustulidae, is redescribed, and its life cycle was resolved experimentally. The prosobranch snail Aylacostoma chloroticum Hylton Scott (Thiaridae), collected in the Yacyretá Dam, Province of Misiones, Argentina, was found naturally infected with cercariae that lacked pigmented eyespots, and possessed 7 pairs of penetration glands, 8 pairs of flame cells, and a V-shaped excretory vesicle. The cercariae developed in oval cysts, which were found on fin rays, vertebrae, and spines of poeciliid and tetragonopterid fish species. Adults were obtained experimentally from Hoplias malabaricus (Erythrinidae) infected with metacercariae from albino Gymnocorymbus ternetzi (Tetragonopteridae), which had been exposed to emerging cercariae. Adults were also found in naturally infected H. malabaricus collected in the Yacyretá Dam. The morphology of the cercariae, and the characteristics of the life cycle show that P. lutzi should be included in the Cryptogonimidae.

Influences of N-Fixing and Non-N-Fixing Vegetation and Invasive Fish on Water Chemistry of Hawaiian Anchialine Ponds

In coastal waters, it remains unclear how terrestrial invasive species might alter nutrient availability and thus affect bottom-up control of primary production. Anchialine ponds are tidal- and groundwater-fed coastal water bodies without surface connections that provide convenient model systems in which to examine terrestrial to aquatic nutrient flow. To investigate how N-fixing and non-N-fixing terrestrial vegetation and invasive consumers affect water quality, we compared dissolved nutrient concentrations and physicochemical parameters among Hawaiian anchialine ponds surrounded by: (1) the nonnitrogen (N)-fixing tree Thespesia populnea (milo), (2) the exotic N-fixing tree Prosopis pallida (kiawe), (3) no tree cover, and (4) no tree cover but containing populations of invasive poeciliid fish. Average (±1 SE) concentration of dissolved inorganic nitrogen (DIN) across all ponds (51.15 µM ±3.1; n = 17) was high, but there was no discernible pattern among pond types. Model results suggest that leaf litter breakdown from the exotic N-fixing tree contributed <0.02% of total dissolved nitrogen (TDN) in the water column at high tide. However, additions of phosphorus (P), especially from litter of non-N-fixing T. populnea, resulted in ratios of DIN to soluble reactive phosphorus (SRP) that varied between 4.5:1 and 35:1 across pond types. Our study suggests that: (1) DIN concentrations were more dependent on watershed-scale N inputs than local tree cover or the presence of poeciliids, (2) low water residence times in these ponds reduce effects of local biota on water nutrient chemistry, and (3) P from anthropogenic inputs and/or biota cause alterations to DIN:SRP ratios in anchialine pond water that may potentially alter primary production rates in these coastal ecosystems.

Sexual Selection: Placentation, Superfetation, and Coercive Copulation

The evolution of placentas in poeciliid fishes is associated with conception of overlapping litters and male mating strategies becoming more coercive. Sperm competition in ovaries of multiply-inseminated females may favor fertilization of immature eggs during ongoing pregnancies.

The expression of pre- and postcopulatory sexually selected traits reflects levels of dietary stress in guppies.

Environmental and ecological conditions can shape the evolution of life history traits in many animals. Among such factors, food or nutrition availability can play an important evolutionary role in moderating an animal's life history traits, particularly sexually selected traits. Here, we test whether diet quantity and/or composition in the form of omega-3 long chain polyunsaturated fatty acids (here termed ‘n3LC’) influence the expression of pre- and postcopulatory traits in the guppy (Poecilia reticulata), a livebearing poeciliid fish. We assigned males haphazardly to one of two experimental diets supplemented with n3LC, and each of these diet treatments was further divided into two diet ‘quantity’ treatments. Our experimental design therefore explored the main and interacting effects of two factors (n3LC content and diet quantity) on the expression of precopulatory (sexual behaviour and sexual ornamentation, including the size, number and spectral properties of colour spots) and postcopulatory (the velocity, viability, number and length of sperm) sexually selected traits. Our study revealed that diet quantity had significant effects on most of the pre- and postcopulatory traits, while n3LC manipulation had a significant effect on sperm traits and in particular on sperm viability. Our analyses also revealed interacting effects of diet quantity and n3LC levels on courtship displays, and the area of orange and iridescent colour spots in the males’ colour patterns. We also confirmed that our dietary manipulations of n3LC resulted in the differential uptake of n3LC in body and testes tissues in the different n3LC groups. This study reveals the effects of diet quantity and n3LC on behavioural, ornamental and ejaculate traits in P. reticulata and underscores the likely role that diet plays in maintaining the high variability in these condition-dependent sexual traits.

Microhabitat use, population densities, and size distributions of sulfur cave-dwelling Poecilia mexicana.

The Cueva del Azufre in Tabasco, Mexico, is a nutrient-rich cave and its inhabitants need to cope with high levels of dissolved hydrogen sulfide and extreme hypoxia. One of the successful colonizers of this cave is the poeciliid fish Poecilia mexicana, which has received considerable attention as a model organism to examine evolutionary adaptations to extreme environmental conditions. Nonetheless, basic ecological data on the endemic cave molly population are still missing; here we aim to provide data on population densities, size class compositions and use of different microhabitats. We found high overall densities in the cave and highest densities at the middle part of the cave with more than 200 individuals per square meter. These sites have lower H2S concentrations compared to the inner parts where most large sulfide sources are located, but they are annually exposed to a religious harvesting ceremony of local Zoque people called La Pesca. We found a marked shift in size/age compositions towards an overabundance of smaller, juvenile fish at those sites. We discuss these findings in relation to several environmental gradients within the cave (i.e., differences in toxicity and lighting conditions), but we also tentatively argue that the annual fish harvest during a religious ceremony (La Pesca) locally diminishes competition (and possibly, cannibalism by large adults), which is followed by a phase of overcompensation of fish densities.

Variation in Melanism and Female Preference in Proximate but Ecologically Distinct Environments

AbstractVariation in pigmentation traits is an effective window to evolutionary processes due to their importance for survival and reproduction. In particular, one of the leading hypotheses for the maintenance of conspicuous pigmentation in natural populations is its signaling function in mate choice. Here, we demonstrate the occurrence of melanism in poeciliid fishes of the genus Poecilia that inhabits toxic, hydrogen sulfide springs in southern Mexico and the absence of melanism from closely related populations in reference habitats lacking hydrogen sulfide. Assays of female mate preference in both habitat types were used to examine whether divergence in female preference for melanism contributes to its maintenance in hydrogen sulfide springs. We found significant variation in female preferences for melanistic males. Specifically, melanistic females from the toxic spring exhibited a significant preference for melanistic males, while non‐melanistic females from the same population exhibited no preference. Females from the non‐sulfidic reference population discriminated strongly against melanistic males. Preferences of melanistic females appear to be a significant force in the maintenance of melanism in sulfidic habitats and suggest a change in preference as the divergence from non‐sulfidic ancestors. Potential polymorphism in preference within the hydrogen sulfide spring indicates that preference for melanistic males may not be environmentally controlled. Thus, a change in preference following divergence can promote the maintenance of variation in pigmentation within populations and between divergent habitats.

Variation in the Visual Habitat May Mediate the Maintenance of Color Polymorphism in a Poeciliid Fish

The conspicuousness of animal signals is influenced by their contrast against the background. As such, signal conspicuousness will tend to vary in nature because habitats are composed of a mosaic of backgrounds. Variation in attractiveness could result in variation in conspecific mate choice and risk of predation, which, in turn, may create opportunities for balancing selection to maintain distinct polymorphisms. We quantified male coloration, the absorbance spectrum of visual pigments and the photic environment of Poecilia parae, a fish species with five distinct male color morphs: a drab (i.e., grey), a striped, and three colorful (i.e., blue, red and yellow) morphs. Then, using physiological models, we assessed how male color patterns can be perceived in their natural visual habitats by conspecific females and a common cichlid predator, Aequidens tetramerus. Our estimates of chromatic and luminance contrasts suggest that the three most colorful morphs were consistently the most conspicuous across all habitats. However, variation in the visual background resulted in variation in which morph was the most conspicuous to females at each locality. Likewise, the most colorful morphs were the most conspicuous morphs to cichlid predators. If females are able to discriminate between conspicuous prospective mates and those preferred males are also more vulnerable to predation, variable visual habitats could influence the direction and strength of natural and sexual selection, thereby allowing for the persistence of color polymorphisms in natural environments.

Testing synaptic plasticity in dynamic mate choice decisions: N-methyl D-aspartate receptor blockade disrupts female preference.

Social behaviours such as mate choice require context-specific responses, often with evolutionary consequences. Increasing evidence indicates that the behavioural plasticity associated with mate choice involves learning. For example, poeciliids show age-dependent changes in female preference functions and express synaptic-plasticity-associated molecular markers during mate choice. Here, we test whether social cognition is necessary for female preference behaviour by blocking the central player in synaptic plasticity, NMDAR (N-methyl d-aspartate receptor), in a poeciliid fish, Xiphophorus nigrensis. After subchronic exposure to NMDAR antagonist MK-801, female preference behaviours towards males were dramatically reduced. Overall activity levels were unaffected, but there was a directional shift from 'social' behaviours towards neutral activity. Multivariate gene expression patterns significantly discriminated between females with normal versus disrupted plasticity processes and correlated with preference behaviours-not general activity. Furthermore, molecular patterns support a distinction between 'preference' (e.g. neuroserpin, neuroligin-3, NMDAR) and 'sociality' (isotocin and vasotocin) gene clusters, highlighting a possible conservation between NMDAR disruption and nonapeptides in modulating behaviour. Our results suggest that mate preference may involve greater social memory processing than overall sociality, and that poeciliid preference functions integrate synaptic-plasticity-oriented 'preference' pathways with overall sociality to invoke dynamic, context-specific responses towards favoured males and away from unfavoured males.

Mating behavior and its morphological correlates in two color morphs of Girardinus metallicus (Pisces: Poeciliidae), a species previously thought not to exhibit courtship display

Girardinus metallicus is a Cuban poeciliid fish whose social behavior has been little studied. The only account involves a colorless morph that is sexually monochromatic and does not exhibit courtship display. We describe the behavior of two other morphs (black and yellow) that exhibit sexual dichromatism. We observed courtship displays in black but not yellow males. Contrary to the pattern in most poeciliids, black males exhibit long gonopodia and courtship; typically, longer gonopodia evolve in species without courtship, because they facilitate coercive mating but circumvent female choice. We focused on the black morph to address whether morphological traits are favored by sexual selection. Larger males with longer gonopodia courted and attempted copulations more often. Black area was not associated with intersexual interactions, but was positively associated with aggressiveness. Dominant males attempted more copulations, consistent with the idea that black coloration may be a badge of status. Black males may possess long gonopodia because the gonopodium itself is a target of female choice. However, there was no difference in gonopodium length between black and yellow males, although the latter do not court. We discuss processes that may maintain the polymorphism and prospects for future studies in this intriguing system.

Superficial neuromasts as proprioceptive receptors in poeciliid fish (1050.11)

The sense of position and movement of the paired and/or unpaired fins (“fin”, hereafter) is critical for executing rapid motor behavior in fishes. However, the location of the proprioceptive receptors involved in proprioception of fin movement is unknown; also unknown is whether or not afferent signals generated during fin movement are encoded to a “central body map” in the brain to reference the location of the fin in space. Here we provide results of work to describe and define the role of a mechanosensory receptor, the superficial neuromast, as a proprioceptive receptor in the male Western Mosquitofish, Gambusia affinis (“Mosquitofish”, hereafter). We sought to clarify whether or not superficial neuromasts associated with the male Mosquitofish sexually dimorphic fin serve as proprioceptive receptors for the proprioception of the fin to perform synchronous‐rapid‐rhythmic (SRR) movements essential for non‐cooperative coitus. In male Mosquitofish, we found that the number and distribution of the superficial neuromasts associated with the sexually dimorphic fin were significantly different (p = 2.6704 e‐05) from those found in female Mosquitofish. Removal of superficial neuromasts from the male resulted in significant impairment of SSR movements. These results suggest that the superficial neuromasts associated with the sexually dimorphic fin in the male Mosquitofish should be considered as a proprioceptive receptor.Grant Funding Source: Research supported by NSF‐1137725

How stable are personalities? A multivariate view of behavioural variation over long and short timescales in the sheepshead swordtail, Xiphophorus birchmanni

Many studies have revealed repeatable (among-individual) variance in behavioural traits consistent with variation in animal personality; however, these studies are often conducted using data collected over single sampling periods, most commonly with short time intervals between observations. Consequently, it is not clear whether population-level patterns of behavioural variation are stable across longer timescales and/or multiple sampling periods or whether individuals maintain consistent ranking of behaviours (and/or personality) over their lifetimes. Here, we address these questions in a captive-bred population of a tropical freshwater poeciliid fish, Xiphophorus birchmanni. Using a multivariate approach, we estimate the among-individual variance-covariance matrix (I), for a set of behavioural traits repeatedly assayed in two different experimental contexts (open-field trials, emergence and exploration trials) over long-term (56 days between observations) and short-term (4-day observation interval) time periods. In both long- and short-term data sets, we find that traits are repeatable and the correlation structure of I is consistent with a latent axis of variation in boldness. While there are some qualitative differences in the way individual traits contribute to boldness and a tendency towards higher repeatabilities in the short-term study, overall, we find that population-level patterns of among-individual behavioural (co)variance to be broadly similar over both time frames. At the individual level, we find evidence that short-term studies can be informative for an individual’s behavioural phenotype over longer (e.g. lifetime) periods. However, statistical support is somewhat mixed and, at least for some observed behaviours, relative rankings of individual performance change significantly between data sets.

Testing the Systematic Position and Relationships ofParacreptotrema heterandriaeWithin the Allocreadiidae Through Partial 28s rRNA Gene Sequences

The genus Paracreptotrema Choudhury, Pérez-Ponce de León, Brooks and Daverdin, 2006 (Digenea), comprises 4 species that parasitize poeciliid and cyprinodontid fishes in Middle America. Based on morphological characters exhibited by adults of the type species (Paracreptotrema blancoi Choudhury, Pérez-Ponce de León, Brooks and Daverdin, 2006 ) the genus was tentatively assigned to Allocreadiidae Looss, 1902. We sequenced domains D1-D3 of the 28S rRNA gene of several specimens of Paracreptotrema heterandriae Salgado-Maldonado, Caspeta-Mandujano and Vázquez, 2012 , and analyzed these in a phylogenetic context along with 10 allocreadiid taxa plus several xiphidiatan and 2 monorchiatan digeneans, in order to test the proposed inclusion of P. heterandriae in Allocreadiidae. Maximum likelihood and Bayesian inference analyses of 28S rRNA gene sequences confirmed the classification of Paracreptotrema as an allocreadiid. In the phylogenetic trees, Paracreptotrema appears to be the sister taxon of Creptotrematina aguirrepequenoi Jiménez-Guzmán, 1973, and both are closely related to Auriculostoma Scholz, Aguirre-Macedo, and Choudhury, 2004, with Wallinia Pearse, 1920, as the sister taxon of all these genera; nevertheless, the interrelationships between these 4 genera were not totally resolved. ITS2 sequences of several specimens of P. heterandriae showed null intraspecific variation. Scanning electron microscopy microphotograph demonstrated the presence of 13 and 4 dome-like papillae arranged around the outer and inner edges of the oral sucker, respectively.

Assessing the Link Between Coastal Urbanization and the Quality of Nekton Habitat in Mangrove Tidal Tributaries

To assess the potential influence of coastal development on habitat quality for estuarine nekton, we characterized body condition and reproduction for common nekton from tidal tributaries classified as undeveloped, industrial, urban or man-made (i.e., mosquito-control ditches). We then evaluated these metrics of nekton performance, along with several abundance-based metrics and community structure from a companion paper (Krebs et al. 2013) to determine which metrics best reflected variation in land-use and in-stream habitat among tributaries. Body condition was not significantly different among undeveloped, industrial, and man-made tidal tributaries for six of nine taxa; however, three of those taxa were in significantly better condition in urban compared to undeveloped tributaries. Palaemonetes shrimp were the only taxon in significantly poorer condition in urban tributaries. For Poecilia latipinna, there was no difference in body condition (length–weight) between undeveloped and urban tributaries, but energetic condition was significantly better in urban tributaries. Reproductive output was reduced for both P. latipinna (i.e., fecundity) and grass shrimp (i.e., very low densities, few ovigerous females) in urban tributaries; however a tradeoff between fecundity and offspring size confounded meaningful interpretation of reproduction among land-use classes for P. latipinna. Reproductive allotment by P. latipinna did not differ significantly among land-use classes. Canonical correspondence analysis differentiated urban and non-urban tributaries based on greater impervious surface, less natural mangrove shoreline, higher frequency of hypoxia and lower, more variable salinities in urban tributaries. These characteristics explained 36 % of the variation in nekton performance, including high densities of poeciliid fishes, greater energetic condition of sailfin mollies, and low densities of several common nekton and economically important taxa from urban tributaries. While variation among tributaries in our study can be largely explained by impervious surface beyond the shorelines of the tributary, variation in nekton metrics among non-urban tributaries was better explained by habitat factors within the tributary and along the shorelines. Our results support the paradigm that urban development in coastal areas has the potential to alter habitat quality in small tidal tributaries as reflected by variation in nekton performance among tributaries from representative land-use classes.

Nekton Community Structure Varies in Response to Coastal Urbanization Near Mangrove Tidal Tributaries

To assess the potential influence of coastal development on estuarine-habitat quality, we characterized land use and the intensity of land development surrounding small tidal tributaries in Tampa Bay. Based on this characterization, we classified tributaries as undeveloped, industrial, urban, or man-made (i.e., mosquito-control ditches). Over one third (37 %) of the tributaries have been heavily developed based on landscape development intensity (LDI) index values >5.0, while fewer than one third (28 %) remain relatively undeveloped (LDI < 3.0). We then examined the nekton community from 11 tributaries in watersheds representing the four defined land-use classes. Whereas mean nekton density was independent of land use, species richness and nekton-community structure were significantly different between urban and non-urban (i.e., undeveloped, industrial, man-made) tributaries. In urban creeks, the community was species-poor and dominated by high densities of poeciliid fishes, Poecilia latipinna and Gambusia holbrooki, while typically dominant estuarine taxa including Menidia spp., Fundulus grandis, and Adinia xenica were in low abundance and palaemonid grass shrimp were nearly absent. Densities of economically important taxa in urban creeks were only half that observed in five of the six undeveloped or industrial creeks, but were similar to those observed in mosquito ditches suggesting that habitat quality in urban and mosquito-ditch tributaries is suboptimal compared to undeveloped tidal creeks. Furthermore, five of nine common taxa were rarely collected in urban creeks. Our results suggest that urban development in coastal areas has the potential to alter the quality of habitat for nekton in small tidal tributaries as reflected by variation in the nekton community.

Retinoic Acid Is Involved in the Metamorphosis of the Anal Fin into an Intromittent Organ, the Gonopodium, in the Green Swordtail (Xiphophorus hellerii)

In poeciliid fish the male anal fin has been transformed into a gonopodium, an intromittent organ required for internal fertilization. Elevated testosterone levels induce metamorphosis of a subset of anal fin rays to grow and form the specialized terminal structures of the gonopodium. The molecular mechanisms underlying these processes are largely unknown. Here, we investigated whether retinoic acid (RA) signaling is involved in gonopodium development in the swordtail Xiphophorus hellerii. We showed that aldh1a2, a RA synthesizing enzyme, and the RA receptors, rar-ga and rar-gb, are expressed in anal fins during metamorphosis. aldh1a2 expression is regulated by testosterone in a concentration-dependent manner and is up-regulated in both hormone-induced and naturally developing gonopodia. Androgen receptor (ar), a putative regulator of gonopodial development, is co-expressed with aldh1a2 and the RA receptors in gonopodial rays. Importantly, experimental increase of RA signaling promoted growth of the gonopodium and increased the number of new segments. Based on gene expression analyses and pharmacological manipulation of gonopodium development, we show that the RA signaling pathway is activated in response to androgen signaling and promotes fin ray growth and development during the metamorphosis of the anal fin into the gonopodium.

Sexual conflict and the function of genitalic claws in guppies (Poecilia reticulata)

Poeciliid fish, freshwater fish with internal fertilization, are known for the diversity of structures on the male intromittent organ, the gonopodium. Prominent among these, in some species, is a pair of claws at its tip. We conducted a manipulative study of these claws in the guppy, Poecilia reticulata, to determine if these aid in transferring sperm to resistant females. We compared the sperm transfer rates of clawed versus surgically declawed males attempting to mate with either receptive or unreceptive (i.e. resistant) females. Our analyses demonstrate that the gonopodial claws function to increase sperm transfer to unreceptive females during uncooperative matings but not during receptive matings. Up to threefold more sperm were transferred to unreceptive females by clawed than declawed males. These data suggest that the claw is a sexually antagonistic trait, functioning to aid in transferring sperm to resistant females, and implicate sexual conflict as a selective force in the diversification of the gonopodium in the Poeciliidae.