Reduction and variability of trunk spines in the acanthocephalan Corynosoma cetaceum: the role of physical constraints on attachment

Abstract. In this study, we investigated a functional trade-off between trunk attachment and trunk-spine development in the acanthocephalan Corynosoma cetaceum . The worms live attached to the stomach and upper intestine of their cetacean definitive hosts, using the proboscis and spiny foretrunk as the main holdfast; the spiny hindtrunk can also attach by bending ventrally. When the hindtrunk bends, ventral compression generates an anterior fold (AF) and a posterior fold (PF). A morphological analysis based on 7,823 individuals collected from 10 franciscana dolphins, Pontoporia blainvillei , revealed that spines were smaller and more variable in size and occurrence in the folds than on neighboring areas; the growth of fold spines seemed to be inhibited to various degrees. Spines were more reduced in the AF than in the PF, and spines of both folds were more reduced in females than in males. Patterns of reduction appeared to be directly related to the intensity of fold compression associated with hindtrunk bending. Fold compression could induce plastic inhibition of spine growth, and/or could make fold spines maladaptive, spines being reduced by natural selection. Apparently, fold spines neither contact the substrate, nor are they exposed to the environment when the hindtrunk attaches. Therefore, fold spines could have reduced, or lost, their primary function, at least in the definitive host. The reduction and variability of spines in C. cetaceum seem to be unique among Corynosoma species.     

Polymorphus arctocephali Smales, 1986, a synonym of Corynosoma cetaceum Johnston & Best, 1942 (Acanthocephala: Polymorphidae)

Polymorphus arctocephali (Acanthocephala: Polymorphidae) was differentiated from P. cetaceum based on patterns in the trunk spine distribution and slight morphometric differences. The comparison of both species involved samples from South Australia and did not include P. cetaceum from South America. In this paper we re-examine the systematic position of P. arctocephali based on a more detailed morphological and geographical analysis. Results indicate that P. arctocephali does not differ in trunk spine distribution with respect to P. cetaceum, and that its morphometric differences can be subsumed under the natural variation found within P. cetaceum populations. Therefore, P. arctocephali becomes a junior synonym of P. cetaceum. P. cetaceum was transferred from Corynosoma to Polymorphus due to the absence of genital spines in both sexes. However, adopting the less restrictive definition of genital spines used by several authors, females of P. cetaceum could be considered as bearing genital spines. Species of the genus Andracantha also have genital spines, but specimens of P. cetaceum possess a continuous field of trunk spines, which precludes the assignment of this species to Andracantha. Other generic level characters, as well as ecological data, support clearly the transference of P. cetaceum back to Corynosoma. Therefore, this species becomes Corynosoma cetaceum Johnston & Best, 1942.     

Presence of genital spines in a male Corynosoma cetaceum Johnston and Best, 1942 (Acanthocephala)

We collected 83 females and 80 males of Corynosoma cetaceum from 2 common dolphins, Delphinus delphis, collected in northern Patagonia (Argentina). Worms were most similar to specimens collected in other South American localities. However, 1 male had 2 spines adjacent to the genital pore and isolated from the rest of body spines. This finding confirms the recent reassignment of C. cetaceum to Corynosoma. Absence of genital spines is suggested to be avoided as the sole criterion to exclude specimens from Corynosoma or Andracantha.     

Assessing host-parasite specificity through coprological analysis: a case study with species of Corynosoma (Acanthocephala: Polymorphidae) from marine mammals

In this paper we report an investigation of the utility of coprological analysis as an alternative technique to study parasite specificity whenever host sampling is problematic; acanthocephalans from marine mammals were used as a model. A total of 252 scats from the South American sea lion, Otaria flavescens, and rectal faeces from 43 franciscanas, Pontoporia blainvillei, from Buenos Aires Province, were examined for acanthocephalans. Specimens of two species, i.e. Corynosoma australe and C. cetaceum, were collected from both host species. In sea lions, 78 out of 145 (37.9%) females of C. australe were gravid and the sex ratio was strongly female-biased. However, none of the 168 females of C. cetaceum collected was gravid and the sex ratio was not female-biased. Conversely, in franciscanas, 14 out of 17 (82.4%) females of C. cetaceum were gravid, but none of 139 females of C. australe was, and the sex ratio of C. cetaceum, but not that of C. australe, was female-biased. In putative non-hosts, the size of worms was similar to that from specimens collected from prey. Results suggest that both acanthocephalans contact sea lions and franciscanas regularly. However, C. australe and C. cetaceum cannot apparently reproduce, nor even grow, in franciscanas and sea lions, respectively. Coprological analysis may represent a useful supplementary method to investigate parasite specificity, particularly when host carcasses are difficult to obtain.     

Sexual dimorphism in Australian Echiniscus (Tardigrada, Echiniscidae) with descriptions of three new species

The presence of males in species of Echiniscus has been reported only as recently as 1987 and is still considered to be rare. However, males have been found to be common in a disproportionately large number of species of this genus found in Australia. Evidence of sexual dimorphism is presented for five species (three of which are new to science), females and males of which are differentiated on the basis of distinctive gonopores. In all five species males are shorter with longer clavae. Males of Echiniscus jamesi sp. nov., Echiniscus rodnae sp. nov. and Echiniscus curiosus sp. nov. have significantly longer claws. In E. rodnae sp. nov. and E. curiosus sp. nov. males have significantly longer lateral trunk appendages and the dorsal trunk appendage combinations of males of the latter are different from those of females. The widespread distribution of bisexual species of Echiniscus in xeric habitats in Australia does not support the current theory that parthenogenetic species are favoured in such environments.     

Genders in Juniperus thurifera have different functional responses to variations in nutrient availability

? Differences in reproductive investment can trigger asymmetric, context-dependent, functional strategies between genders in dioecious species. However, little is known about the gender responses of dioecious species to nutrient availability. ? We experimentally fertirrigated a set of male and female Juniperus thurifera trees monthly for 2 yr. Water potential, photosynthesis rate and stomatal conductance were measured monthly for 2 yr, while shoot nitrogen (N) concentration, carbon isotopic composition (δ(13) C), branch growth, trunk radial growth and reproductive investment per branch were measured yearly. ? Control males had lower gas exchange rates and radial growth but greater reproductive investment and higher water use efficiency (WUE; as inferred from more positive δ(13) C values) than females. Fertirrigation did not affect water potential or WUE but genders responded differently to increased nutrient availability. The two genders similarly increased shoot N concentration when fertilized. The increase in shoot N was associated with increased photosynthesis in males but not in females, which presented consistently high photosynthetic rates across treatments. ? Our results suggest that genders invest N surplus in different functions, with females presenting a long-term strategy by increasing N storage to compensate for massive reproductive masting events, while males seem to be more reactive to current nutrient availability, promoting gas-exchange capacity.     

Allocation to floral structures in Thalictrum pubescens (Ranunculaceae), a cryptically dioecious species

Females of Thalictrum pubescens produce stamens that contain sterile pollen, whereas males are both functionally and morphologically unisexual. This study examines the investment in stamen production by females of T. pubescens by comparing the female structures with those of their fully functional male counterparts. Stamens from females had the same biomass and contained the same amount of nitrogen and phosphorus as stamens from males. Anther size was the same in males and females, but filaments were longer in stamens from males. Females produced more pollen per anther than males, and pollen size was the same in both sexes. Within flowers, there was a positive correlation between the amount of pollen per anther and the length of anthers in males, but not in females. This would be expected if males growing in better environmental conditions or with greater vigour invested more resources in pollen production, thereby increasing fitness. Females, who receive no fitness benefits from increased pollen production, did not show this pattern. There was also evidence of a trade-off within female flowers between the number of stamens and the number of pistils. This trade-off was noted in conditions when variance among plants was reduced, namely in the field during a year when flower size was particularly small and in a previous glasshouse study. Therefore, it appears that when environmental variance is low, stamens are produced at the expense of producing more pistils, and hence seeds. In conclusion, stamen production does not appear to be inconsequential to females of Thalictrum pubescens.     

Sexually dimorphic legs in a neotropical harvestman (Arachnida,Opiliones): Ornament or weapon?

The evolution of sexually dimorphic traits has been the focus of much theoretical work, but empirical approaches to this topic have not been equally prolific. Males of the neotropical family Gonyleptidae usually present a strong fourth pair of legs armed with spines, but their functional significance is unknown. We investigated the putative functions of the leg armature in the harvestman Neosadocus maximus. Being a non-visual species, the spines on male legs can only be perceived by females through physical contact. Thus, we could expect females to touch the armature on the legs of their mates if they were to evaluate it. However, we found no support for this hypothesis. We did show that (1) leg armature is used as a weapon in contests between males and (2) spines and associated sensilla are sexually dimorphic structures involved in "nipping behavior", during which a winner emerged in most fights. Finally, we demonstrate that five body structures directly involved in male-male fights show positive allometry in males, presenting slopes higher than 1, whereas the same structures show either no or negative allometry in the case of females. In conclusion, leg armature in male harvestmen is clearly used as a device in intrasexual contests.     

An Extreme Case of Interspecific Territoriality: Male Anthidium manicatum (Hymenoptera,Megachilidae) Wound and Kill Intruders

Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.     

Spine production is induced by fire: a natural experiment with three Berberis species

Earlier studies indicate that some plant species allocate more mass to produce longer spines in shoots resprouting after browsing. Here we present, for the first time, evidence that fire induces a similar response. Many terrestrial herbivores may benefit from fire through the enhanced availability of fast growing species colonizing or re-sprouting in burned areas. It is less clear whether post-fire plant growth responds to the enhanced risk of herbivory by an increased investment in defensive traits. In this study, we tested whether the production of spines is influenced by the set of environmental conditions that result from fire events. We compared the resource allocation pattern of resprouting shoots from three Berberis species growing in two areas that burned 1999 with samples collected from unburned areas within the same plant communities. We divided the shoot into three main components: supporting tissue (twigs), assimilating tissue (leaves) and defensive structures (spines). We found that plants resprouting after fire allocated more mass to spines and leaves but not twigs. This resulted in a higher density of both spines and leaves. Spines were significantly longer in plants resprouting after fire. Leaves were shorter at the apical end of the shoot, but did not show any significant change in size following fire. We suggest that this type of post-fire response may be a general adaptation to pruning and leaf picking by browsing herbivores in arid and semi-arid regions. Changes in the browsing pressure following fire will determine the fitness value of this response.     

Investment in Mate Guarding May Compensate for Constraints on Ejaculate Production in the Cricket Gryllodes sigillatus

Although there is a corpus of evidence that females of many taxa are choosy about males, there is less information on how males may react to females of different 'quality' (i.e. potential fecundity). The cricket Gryllodes sigillatus shows distinct mate guarding behaviour. We examined how long males mate guard females of different sizes (reflecting egg load and potential fecundity). We also examined the sperm number in ampullae donated to females of different sizes to see if males make a concomitant difference in investment in ejaculate. We also examined mate-guarding behaviour and ejaculate size of males mated to virgin and nonvirgin females of the same size to see if males equate size with increased age and increased likelihood of mating (increased sperm competition). The results showed that males mate guard larger females for longer but make no difference in ejaculate investment between sizes of female. Males make no significant difference in mate guarding investment or ejaculate investment between virgins and nonvirgins of the same size. There is evidence that other species of crickets do alter their ejaculate depending on the female size and mating history, but have less distinct guarding behaviour. We suggest that mate-guarding investment in G. sigillatus may serve a similar function to that of ejaculate investment in other crickets.     

Adult Responses to Larval Population Size in the Almond Moth, Cadra cautella

Juvenile population size may affect the potential for future mating opportunities and therefore potentially sperm competition; this may favour ontogenetic adjustments in sperm production. Theory predicts that males should optimize their ejaculatory investment in accordance with the risk of sperm competition. Evidence for these theories is typically revealed in males of highly polyandrous species. Whether such responses to environmental cues exist for females, or are maintained in mildly polyandrous species in which most females do not re-mate, is unknown. Male lepidopterans produce normal, fertilizing sperm (eupyrene) and non-fertilizing (apyrene) sperm. Apyrene sperm are associated with reduced female receptivity, suggesting a role in sperm competition. We tested the effect of juvenile population size on life-history parameters and reproductive investment in the mildly polyandrous almond moth, Cadra cautella , a species in which current male ejaculate traits suggest previous selection for paternity protection consistent with a sperm-competitive environment. Larvae were reared at high (H) or low population sizes (L). We recorded larval development time, adult longevity and male gametic investment. Our results show a response by adults to signals in the juvenile environment. H males transferred more apyrene, but not eupyrene sperm. We also examined potential trade-offs between somatic characters and reproductive behaviours. Larval duration was longer for H individuals, females and heavier individuals. Further, H females and L males lived longer than L females. Our data are consistent with the theory that males should adjust their reproductive investment in accordance with sperm competition risk.     

The effects of copulation duration in the bruchid beetle Callosobruchus maculatus

Control over copulation duration is a potentially importantgenerator of sexual conflict that has received little empiricalattention. The copulatory behavior of the bruchid beetle Callosobruchusmaculatus may reflect a sexual conflict over copulation duration.Males have spines on their intromittent organs that puncturethe female reproductive tract, and females kick their matesduring copulation. If females are prevented from kicking, copulationslast longer and the injuries females sustain are more severe.Males supposedly use the spines as anchors to prolong copulationduration, and females kick to terminate copulations. We manipulatedcopulation duration experimentally and quantified its effectson male and female fitness components to test whether or notthere is a conflict over copulation duration in C. maculatus.Females did not suffer from long copulations but instead experiencedincreased lifetime fecundity. Ejaculate size increased withcopulation duration, and females apparently derive materialbenefits from the ejaculates. Males that mated first and hadlong copulations were relatively unsuccessful when competingwith sperm from other males. However, there was a trend forfemale remating propensity to decrease with long copulationdurations, and first males may therefore also benefit from longcopulations. The copulation duration of the second male to matedid not have a significant effect on sperm precedence. We concludethat even though it seems likely that the male spines have evolvedto act as an anchor during copulation, there seems to be littleconflict over copulation duration per se in C. maculatus.     

Molecular phylogeny of Corynosoma Lühe, 1904 (Acanthocephala), based on 5.8S and internal transcribed spacer sequences

Species of Corynosoma (Acanthocephala) are distributed worldwide as parasites of marine mammals and sea birds. Species diagnosis is based on morphological characters, including the size and number of hooks in the proboscis and the number of spines in the dorsal and ventral regions of the body. We inferred the phylogenetic relationships of 10 nominal species of Corynosoma through analysis of internal transcribed spacers (ITS-1, ITS-2) and 5.8S ribosomal RNA sequences. Nucleotide distances between species of Corynosoma ranged from 0.4 to 11% for ITS sequences. Maximum parsimony and likelihood analyses indicated that species of Corynosoma that inhabit hosts in the marine environment form a monophyletic assemblage, but yielded conflicting hypotheses for the relationship of Corynosoma cetaceum to other members of the genus. However, parsimony and likelihood analyses were consistent for many Corynosoma sister species relationships of (e.g., C. australe plus C. bullosum, C. validum plus C. villosum, C. caspicum plus C. magdaleni, and C. enhydri plus C. strumosum). This phylogenetic framework was used to evaluate taxonomic controversies concerning C. cetaceum and C. caspicum.     

Season and testosterone affect contractile properties of fast calling muscles in the gray tree frog Hyla chrysoscelis

In anurans, circulating levels of androgens influence certain secondary sexual characteristics that are expressed only during the breeding season. We studied the contractile properties of external oblique muscles (used to power sound production) in a species of North American gray tree frog, Hyla chrysoscelis, during the breeding season and also in testosterone-treated captive males and females after the breeding season. Compared with the muscles of breeding-season males, the trunk muscles of postbreeding-season males have 50% less mass, 60% longer twitches, and 40% slower shortening velocities. Testosterone levels similar to those found in breeding-season male hylid frogs restore the contractile speed and mass of male trunk muscles and also convert the small slow trunk muscles of females into larger fast-contracting muscles. We conclude that androgens likely play a key role in altering the contractile properties of these muscles in males during the annual cycle, allowing them to operate in the breeding season at the frequencies required to produce the characteristic rapidly pulsed calls of this species. Females as well as nonbreeding-season males do not produce advertising calls, and therefore the slower muscles found in these animals may allow more economic operation of these muscles. The effects of testosterone on female trunk muscles indicate the potential of this hormone in contributing to the sexual dimorphism in size and contractile properties of these muscles, but this dimorphism is likely due to the interaction of more than one hormone.     

Sexual comparisons in immune ability, survival and parasite intensity in two damselfly species

Recent evolutionary studies have suggested that females have a more robust immune system than males. Using two damselfly species (Hetaerina americana and Argia tezpi), we tested if females produced higher immune responses (as phenoloxidase and hydrolytic enzymes), had a higher survival (using a nylon implant inserted in the abdomen and measuring survival after 24h) and fewer parasites (gregarines and water mites) than males. We also tested whether immune differences should emerge in different body areas (thorax vs. abdomen) within each sex with the prediction that only females will differ with the abdomen having a higher immune response than their thorax since the former area, for ecological and physiological reasons, may be a target zone for increased immune investment. Animals were adults of approximately the same age. In both species, females were more immunocompetent than males, but only in H. americana females were immune responses greater in the abdomen than in the thorax. However, there were no differences in survival and parasite intensity or the probability of being parasitised between the sexes in either of the two species. Thus, this study lends partial support to the principle that females are better at defending than males despite the null difference in parasitism and survival.     

Does heterospecific seminal fluid reduce fecundity in interspecific copulation between seed beetles?

Reproductive interference through mating between related species can cause fitness reduction and affect population dynamics of the interacting species. In experimental matings between two seed beetles, Callosobruchus chinensis and Callosobruchus maculatus, C. maculatus females, but not C. chinensis females, suffer from significant loss of fecundity when conspecific mating is followed by heterospecific mating. We hypothesized that male traits associated with sexual conflict, which are often harmful to females, pleiotropically affect fitness of heterospecific females through interspecific mating. We examined the effect of ejaculate of C. chinensis males on C. maculatus females as the cause of the fecundity loss in C. maculatus females due to interspecific copulation. We found that frequent interspecific copulation occurred between C. maculatus females and C. chinensis males, but not between C. chinensis females and C. maculatus males, resulting in frequent interspecific ejaculate transfer from C. chinensis males to C. maculatus females. However, injection of the extract from C. chinensis male reproductive organs into C. maculatus females did not significantly affect C. maculatus fecundity compared with saline injection, indicating that the effect of the heterospecific ejaculate transfer on fecundity is negligible. We suggest that other harmful male traits such as genital spines of C. chinensis males are mainly responsible for the fecundity reduction in C. maculatus females that have experienced interspecific mating.     

Mating behavior in eight rotifer species: using cross-mating tests to study species boundaries

We characterized the mating behavior of eight freshwater rotifer species from five different families of Monogononts (Asplanchnidae, Brachionidae, Euchlanidae, Lecanidae, and Trichocercidae). Were corded the percentages of male-female encounters resulting in mating attempts and the number of completed copulations. The characterization of mating behavior also included data on: (1) duration of copulation, (2) female and male swimming speed, (3) sites of initial mating attempts, (4) sites of copulation, and (5) sizes of males and females. In the five families investigated, cross-mating tests may be used to assess the ability of males to discriminate conspecifics, establishing limits to gene flow. We also present data on two families, Brachionidae and Lecanidae where we used cross-mating tests to compare discrimination between conspecific females and those of other species. Brachionus calyciflorus males attempted mating with females of the closely related genus Plationus, but did not attempt mating with Keratella Americana females. Males of other three Brachionidae species only attempted mating with nonspecific females. Lecane quadridentata males attempted to mate with females of two different Lecane species besides their own. Percentages of mating attempts and completed copulations were quantified in all crosses. Lower percentages of copulation after a mating attempt were found among brachionids than among non-brachionids. These data demonstrate that male discrimination of females is an effective pre-mating reproductive barrier in several species of Monogonont rotifers. These results, further suggest that investigations of mating behavior are useful for determining species boundaries in rotifers and perhaps other zooplankters. This revised version was published online in August 2006 with corrections to the Cover Date.     

<Emphasis Type="Italic">Condyloderes kurilensis</Emphasis> sp. nov. (Kinorhyncha: Cyclorhagida)—a new deep water species from the abyssal plain near the Kuril-Kamchatka Trench

A new species of centroderid kinorhynchs, Condyloderes kurilensis sp. nov., collected from muddy sediment at the abyssal plain near the Kuril-Kamchatka Trench, North-West Pacific, is described and illustrated using light and electron microscopy. Previously, members of the genus Condyloderes were found only in nearly shallow water enviroments—C. multispinosus (North Sea), C. paradoxus (Bengal Bay), C. setoensis (Tanabe Bay, Japan), C. storchi (Argentina coast), and C. megastigma (Korea Strait). The new species is characterized by the presence of lateroventral cuspidate spines only on trunk segment 8 in both sexes, paired ventromedial appendages (tubules) on trunk segments 7 and 8 in females only, and dorsolateral spines on trunk segment 10 in males only. Condyloderes kurilensis sp. nov. constitutes the sixth species of the genus Condyloderes described so far and the only second abyssal species of the Kinorhyncha described from the North-West Pacific. A key to the six species of the genus Condyloderes is proposed.