Parasitism of Leptasterias spp. (Echinodermata: Asteroidea) by the ciliated protozoan Orchitophrya stellarum (Scuticociliata)

Abstract. The parasitic ciliate Orchitophrya stellarum was found in the testes of brooding, winter-spawning Leptasterias spp. from San Juan Island, Washington, but not in the testes of Leptasterias spp. from the Lynn Canal, Alaska. Dense populations of the ciliate were localized within the fuller areas of testes, where sperm counts were significantly reduced. The ciliates were loaded with phagosomes, some of which contained sperms in various stages of digestion. Leptasterias spp. are not as severely impacted by this ciliated protozoan parasite as Pisaster ochraceus. Leptasterias spp. may serve as seasonal hosts for O. stellarum ; this ciliate lives in seawater and in ripe males of winter-brooding and spring-summer broadcasting sea stars. The wet weight of parasitized and non-parasitized males did not differ. The sex ratio did not deviate from the expected (1:1) in any population except at Point Louisa, Alaska, where males outnumbered females. Testis indexes of parasitized males from 2 of the 4 locations on San Juan Island, Washington were significantly reduced relative to non-parasitized males, indicating a loss of sperm output during spawning.     

Evolution of pelagic direct development in the starfish Pteraster tesselatus (Asteroidea: Velatida)

Despite a diversity of larval forms, remarkably conservative features of asteroid development define a larval body plan that occurs throughout the class. However, recent work on the starfish Pteraster tesselatus has documented a highly derived pattern of development. Several features, including radial symmetry, parallel embryonic and adult axes of symmetry, absence of a preoral lobe, and formation of coeloms in the adult orientation from seven separate enterocoels, have not been reported in asteroids before. The complete absence of the larval body plan features that are found in other asteroids, indicates that P. tesselatus develops directly from the embryo to the juvenile and has a pelagic, nonfeeding (lecithotrophic), but nonlarval mode of development. I postulate that direct development evolved over an extended period in a lineage of brooding, deep-sea velatid (probably pterastcrid) ancestors of P. tesselatus. Selection for increased developmental efficiency (loss of nonfunctional larval features) in the brooded offspring, could explain the lack of larval settlement structures, the nonlarval arrangement of coeloms, the lack of a preoral lobe, the transverse orientation of the juvenile disc, and the lack of bilateral symmetry. The pattern of coclomogenesis could have been derived from that of other velatids (e.g. solasterids) by relatively simple changes in timing and orientation of entcroeoel formation. Rotation and posterior translation of the coelomic fate map of the archenteron prior to enlerocoel formation would produce the coelomic compartments in the adult orientation that characterizes direct development in P. tesselatus. These unusual developmental features lead to a radically different interpretation for the evolution of the pelagic ‘larva’ of P. tesselatus: (1) evolution of benthie brooding, (2) extreme simplification of development involving the loss of all larval features from the life cycle, and (3) subsequent re-evolution of pelagic development. In the case of P. tesselatus, where all larval structures were lost, there do not seem to be functional constraints preventing the re-evolution of pelagic development. Analysis of pelagic and benthie larvae, in other asteroids, suggests that major ecological transitions in life histories need not be associated with substantia] changes in morphology. The loss of pelagic development should have occurred repeatedly and should be readily reversible. These findings have interesting implications for the loss and evolution of pelagic dispersal in the life histories of marine benthie invertebrates.     

Effects of salinity and feeding on arm regeneration in the starfish Luidia clathrata (Say, 1825) (Echinodermata: Asteroidea)

The aim of this study was to determine: (1) salinity that maximizes arm regeneration in the starfish Luidia clathrata; and (2) if low food consumption or low salinity is the cause of the reduced rate of arm regeneration seen at low salinities.. An estimated salinity of 33 g kg^?1 produced maximum regeneration of two arms in L. clathrata. This salinity is typically greater than that found in bays that are the usual habitat for the species. Equivalent food consumption by L. clathrata in salinities of 20 g kg^?1 and 30 g kg^?1 resulted in greater regeneration in salinity of 30 g kg^?1, suggesting lower food consumption at low salinities is not the primary reason for lower regeneration. Higher food consumption compared to lower food consumption, both tested in salinity of 30 g kg^?1, did not increase regeneration, but resulted in more storage of energy in the pyloric caeca.     

Phylogeny and reproduction in Syllidae (Polychaeta)

The phylogeny of 12 representatives of the four syllid subfamilies is analysed using 44 morphological characters and 96 character states. The analysis is designed to answer two questions: are the four subfamilies monophyletic, and how are the different reproductive strategies related? Syllids show two modes of reproduction, epigamy and schizogamy. Epigamy is mainly constrained to two of the four subfamilies, Eusyllinae and Exogoninae but occurs also in some members of Autolytinae. Schizogamy can be of two types–scissiparity and gemmiparity–and both types are found in Syllinae and Autolytinae. Several modes of brood protection also occur in the family; in some epigamous species the eggs and even juveniles are attached to the main animal, whereas in brooding schizogamous species the stolons care for the young. Brood protection appears in Exogoninae, a few species of Eusyllinae, and some Autolytinae. The resulting tree indicates that members exhibiting brood protection are constrained to one clade, and that it is uncertain whether epigamy or schizogamy is the plesiomorphic reproductive mode. Three of the four subfamilies, Autolytinae, Exogoninae and Syllinae, are supported in the analysis whereas Eusyllinae is found to be polyphyletic.     

Photoperiodic regulation of feeding and reproduction in a brooding sea star from central California

Summary

Individuals of the sea star genus Leptasterias from the Monterey Peninsula, California, were maintained for 34 months under two contrasting regimes of seasonally changing photoperiod. In one regime the lights turned on and off in phase with local sunrise and sunset; in the other the daily photoperiod was kept 6 months out of phase with ambient photoperiods. Annual cycles of feeding, gametogenesis, and embryo brooding of the out-of-phase animals were all shifted out of phase with respect to those of the in-phase animals. These small, relatively hardy sea stars appear well suited for more detailed analyses of photoperiodism in asteroids.     

Food intake in Blattella germanica (L.) nymphs affects hydrocarbon synthesis and its allocation in adults between epicuticle and reproduction

The causal relationship between food intake and hydrocarbon synthesis was examined in vivo and in vitro. Fed Blattella germanica (L.) nymphs synthesized hydrocarbons in a stage‐specific manner, with high rates occurring in the first 6 days of a 13‐day last stadium, in relation to feeding. A similar pattern was exhibited in vitro by sternites and tergites from fed nymphs. In contrast, starved nymphs synthesized hydrocarbons at normal rates for the first 2 days, but then synthesis declined and ceased by day 6. Their abdominal sternites and tergites displayed a similar biosynthetic pattern in vitro, showing that starved tissues lost the capacity to synthesize hydrocarbons, even when provided appropriate nutrients. Synthesis resumed within 2 days of being fed on day 6, reaching a maximum rate 6 days later. Some hydrocarbon appeared on the nymphal cuticle, but almost 4‐fold more hydrocarbon was internal in hemolymph lipophorin, fat body, and the developing imaginal cuticle. Because most hydrocarbon synthesized in nymphs provisions the adult, and synthesis is related to food intake, we examined trade‐offs in allocations in food‐limited insects. Nymphs provided with insufficient quantities of food allocated normal amounts of hydrocarbons to the nymphal epicuticle, but molted into smaller adults with significantly less internal hydrocarbons. These cockroaches directed nearly normal amounts of hydrocarbons to their epicuticle, oocytes, and oothecae, at the cost of internal hydrocarbon reserves for repair and subsequent gonotrophic cycles. Hydrocarbons, thus, appear to serve an important cross‐stadial resource and the object of competition among several nymphal and adult tissues. Arch. Insect Biochem. Physiol. 41:214–224, 1999. © 1999 Wiley‐Liss, Inc.     

Biennial reproduction with embryonic diapause in Lopholithodes foraminatus (Anomura: Lithodidae) from British Columbia waters

Abstract. A paucity of data on the reproductive cycle of crabs in the family Lithodidae inhibits both the development of management strategies and the formulation of hypotheses regarding the evolution of lithodid life histories. Life‐history parameters of Lopholithodes foraminatus from British Columbia, Canada, were investigated based on 26 females maintained in the laboratory and supplementary observations on other living and preserved animals. The rate of embryonic development was determined by measuring the percentage area occupied by yolk in lateral views of eggs removed from brooding females throughout development. Females of L. foraminatus exhibited biennial reproduction including an 18‐month brooding period. Females molted, mated, and extruded eggs in mid‐summer, and did not release larvae until late winter or early spring of the second year after fertilization. Embryogenesis included a 12‐month diapause at the gastrula stage. Females released larvae over a mean interval of 69 d, the longest reported for any lithodid. While the development stage of embryos was observed to be heterogeneous within a brood, no spatial gradient in development rate was observed, calling into question the oxygen limitation hypothesis of extended hatching. Biennial reproduction of individuals of L. foraminatus may be a consequence of a relatively low‐quality habitat. Relative to annual reproduction, biennial reproduction halves the potential rate of increase of a population and increases vulnerability to overharvesting, suggesting that L. foraminatus is not a good candidate for commercial exploitation. The adaptive value of embryonic diapause is uncertain and warrants further research.     

Laboratory observations on the feeding behaviour, reproduction and morphology of Galeodea echinophora (Gastropoda: Gassidae)

Galeodea echinophora fed on Echinocardium cordatum in an aquarium. Every few days, each G. echinophora emerged from the sand and foraged for 1–3 h. On detecting an Echinocardium , the Galeodea stopped locomotion and attacked the buried prey from the surface of the sand. The proboscis was extended down to the prey and a small area of test was cleared of spines. A disc was cut out of the softened test, leaving a hole of about 2 mm in diameter. All flesh except the gut was removed from the prey, the entire procedure taking 50–180 min.
One Galeodea echinophora laid 120 egg capsules on the side of the aquarium. These were kept at 13^oC and hatched after 112–159 days. Within the capsule, veligers fed on the yolk cells of abortive embryos. Juveniles hatched at the crawling stage, and at first secreted strings of mucus acting as drogues, but after 1 day the juveniles crawled over the substratum. They readily attached themselves to adult Echinocardium cordatum , apparently feeding on the epithelium between the spines or tube feet. Dissection revealed entry of the acinar glands into the proboscis gland ducts, a widening of these ducts as they empty into the buccal cavity and a dorsal fold in the anterior oesophagus. Radular teeth and jaws of Galeodea differ little among species; a taxonomic review of the genus is needed.     

Seasonal reproduction and sexual maturity in Odontocymbiola magellanica (Neogastropoda, Volutidae)

Abstract. Members of Odontocymbiola magellanica (Volutidae, Odontocymbiolinae) inhabit Argentinean waters from 35°S to the Magallanes Strait. The reproductive biology of this potential fishery resource has not been studied before. Male individuals reach maturity at 80 mm and females at 90 mm, corresponding to ∼7 and 8 years of age, respectively. Mating was observed in the field in all seasons and females possess live spermatozoa in the bursa copulatrix all year round, suggesting that living sperm can be preserved for several months. Major oviposition events occurred July (winter) to December (summer) during the 5 years sampled, as characterized by aggregation of females coincident with increasing photoperiod from winter to summer solstices, and not, apparently, by increase in water temperature. Males showed gametogenic activity throughout the year. Gamete resorption occurred in both sexes. Ultrastructural studies showed that oocytes are attached to companion cells, which enables cell–cell communication and provides support until evacuation; males showed normal sperm cells and two types of parasperms. If populations of O. magellanica are commercially exploited in the future, we propose a minimum catch size of 100 mm, and a ban on the collection of females and egg capsules during the reproductive season.     

An articulated titanosaur from Patagonia (Argentina): New evidence of neosauropod pedal evolution

Most titanosaur dinosaurs are represented by incomplete skeletal elements lacking articulated pes. An exceptionally preserved specimen from the Late Campanian–Early Maastrichtian strata of Patagonia (Argentina) provides new data on pedal morphology and the evolutionary trends of these huge dinosaurs. This finding is one of the few articulated titanosaur pes known in the world, and shows a phalangeal formula of 2-2-2-2-0. The first three digits possess sickle-shaped claws and the articular facets of ungual phalanges, suggesting mobility in horizontal and vertical planes. A comparative analysis of available record suggests that titanosaurs had a progressive reduction of size and number of pedal phalanges in digits III and IV during the Late Cretaceous.     

Same species,different population dynamics: Spatio-temporal differences of Undaria pinnatifida (Ochrophyta,Phaeophyceae) in the intertidal of North Patagonia,Argentina

Population dynamics can be influenced by physical and biological factors, particularly in stressful environments. Introduced species usually have great physiological plasticity, resulting in populations with different traits. Undaria pinnatifida, a macroalga originally described from northeast Asia, was introduced in Northern Patagonia, Argentina (San Matías Gulf) around 2010. To describe the spatio-temporal variability in population structure and morphometry of U. pinnatifida, we conducted monthly field samplings for 2 years at the intertidal area of two contrasting sites in the San Matías Gulf. Individuals of U. pinnatifida were classified by developmental stage, and their morpho-gravimetric variables were measured. In both intertidal sites juveniles were found in higher proportion during austral autumn and grew and matured during the autumn-winter months (from May onwards), and individuals senesced during early austral summer (December and January). Conversely, density and biomass were largely different between sites, and individuals showed slight morphological variability between sites. Environmental (e.g., nutrient concentration, available substrate) and biological factors (e.g., facilitation, competition) may explain the observed differences. Since there is not a macroalga with U. pinnatifida morphometrical characteristics in the intertidal environments of San Matías Gulf, studying this recent introduction gives us a better understanding of its potential ecological effects.     

Avoidance and escape responses of the sub-antarctic limpet Nacella edgari (Powell) (Mollusca: Gastropoda) to the sea star Anasterias perrieri (Smith) (Echinodermata: Asteroidea)

Summary The sub-antarctic limpet Nacella edgari (Powell) (Patellidae) has avoidance and escape responses to the sea star Anasterias perrieri (Smith) (Asteriidae) in a kelp forest and in the laboratory. Nacella edgari was preyed upon in the field by A. perrieri, but frequency of capture was low, suggesting the effectiveness of defensive behaviors. Both avoidance and escape responses by the limpet consisted of extension and waving of the pallial tentacles, mushrooming and rotation of the shell and flight. However, escape responses were characterized by greatly intensified rotation of the shell through a horizontal plane. This study extends the evaluation of defensive responses in gastropods to a species which occurs in the southern Indian Ocean.     

The feeding behaviour, activity and trappability of wild female Glossina pallidipe in relation to their pregnancy cycle

Female Glossina pallidipes Austen trapped with baited NG2B traps were subjected both to detailed ovarian dissection and to nutritional analysis. Using a calibration curve derived from dissected wild-caught, laboratory-held flies, the field females were assigned by discriminant analysis to each day of the pregnancy cycle. Field females were most available to NG2B traps while carrying the first instar larva. The nutritional characteristics of trapped field females over the pregnancy cycle lead to the following main conclusions. (i) Fat levels increase most rapidly during the egg in utero stage, while CRDW increases significantly only during the larval stages, culminating in a 4 mg increase during the last day of the third larval instar. (ii) The haematin content of the flies indicates that females feed at approximately 3-day intervals and may feed on any day of the pregnancy cycle. (iii) The estimated time of feeding during the day corresponds with the observed time of peak activity, both of which are earlier in the day later in the pregnancy cycle. (iv) The rate of fat usage reveals significantly greater flight activity on day 5 of the cycle than on other days, agreeing with the high trappability on this day, and overall females appear to use fat at twice the rate of males. Reproductive data provide a much more accurate picture of the relative sampling efficiency than do nutritional data, although the latter reveal the general trends correctly.     

Intertidal and Subtidal Fouling Assemblages in a Patagonian Harbour (Argentina, Southwest Atlantic)

The composition of the early stages of intertidal and subtidal fouling assemblages in Comodoro Rivadavia harbour (Argentina, 45°52′ S, 67°28′ W) and the influence of shore level and season on their structure were analysed. At the beginning of each season, stones were glued to the substratum with epoxy putty and distributed along 4 vertical transects at intervals of 20 m, at 3 levels: upper intertidal, middle intertidal, and subtidal. Substrata remained in the field for 84–100 days. A total of 48 samples (4 seasons × 3 levels × 4 replicates) were analysed. Species richness increased with depth, with 6 taxa in the upper intertidal, 23 in the middle intertidal and 31 in the subtidal. Seasonal differences in richness were less distinct. Green, red and brown algae were the dominant groups. Invertebrates were mainly represented by filter-feeding, sessile organisms, such as cheilostome bryozoans, spirorbid polychaetes and acorn barnacles. The barnacle Balanus glandula and the bryozoan Cryptosula pallasiana were the only non-indigenous species found in this study. Highly significant differences in structure among shore levels and seasons were evidenced by a two-way ANOSIM test. The upper intertidal is characterized by the filamentous green algae Urospora penicilliformis and Ulothrix flacca. The barnacle Balanus glandula is the most abundant species in the middle intertidal. The subtidal is defined mainly by the presence of the spirorbid polychaetes Paralaeospira levinseni and Romanchella perrieri, and the keyhole limpet Fissurella radiosa. Ordination of samples by season was less clear than by shore level.     

Community assembly and inventory efficiency at a regional scale: the case of terrestrial Heteroptera (Insecta) in northern Patagonia,Argentina

The first step to understanding the species present in a particular area is to perform inventory and assemblage studies. To obtain a species inventory, it is important to determine parameters such as species richness and relative abundance. This information can be useful for future studies and decision-making purposes in the conservation area. Despite the fundamental role of the terrestrial Heteroptera in ecosystems, they remain poorly known. We expected that the terrestrial Heteroptera species assemblage would be strongly associated with plant communities. Presently, 840 samples were collected in northern Argentine Patagonia during two years, 2013 and 2014. A total of 1950 adults of terrestrial Heteroptera belonging to 12 families, 32 species, and 8 morphospecies were found. Various statistical techniques were applied to correct the observed data for undersampling bias. These suggested that the lower boundary of the summer Heteroptera species richness in northern Patagonia was about 44–54 species. We concluded that the high regional habitat heterogeneity along the west-to-east and south-to-north gradients was paralleled by the turnover of Heteroptera. However, the Patagonian steppe shared a high number of species with the Monte and Subantarctic provinces. The suction sampling technique is an efficient technique to collect Heteroptera in environments with different plant structure and should thus be used in a complementary way with the sweeping technique.     

A new genus and species of darner dragonfl y (Aeshnidae: Odonata) from the lower Eocene of Laguna del Hunco,Patagonia, Argentina

A new genus of Aeshnidae, Huncoaeshna n. gen., based on Huncoaeshna corrugata n. gen., n. sp., is erected from Laguna del Hunco (Ypresian) in Patagonia Argentina. The specimen presents a special kind of preservation with the middle part of the wing wrinkled. The presence of only two fossil specimens of Aeshnidae in South America is surely due to the lack of paleontomologists and collections of fossil insects in the subcontinent.     

A histological study of microsporogenesis in Tarenna gracilipes (Rubiaceae)

We studied the microsporogenesis in Tarenna gracilipes (Hayata) Ohwi, with special attention to the mode of exine deposition and tapetum development. We based this research on light (LM), scanning electron (SEM) and transmission electron microscopic (TEM) observations of developing anthers of T. gracilipes, from the microspore mother cell stage towards anther dehiscence. Evidence is supplied that the microsporogenesis in T. gracilipes can be considered as simultaneous. Columellae, foot layer and tectum develop in a fibrillar matrix. Similar with earlier studies in Rubiaceae species, a single white line formed near the plasmalemma in the extra‐apertural region. The developing endexine dilated into several white line centered lamellae at the apertures. An annulus is formed around the inner surface of each pore. In the mature intine two strata can be distinguished. At the apertures thick onci are formed protruding through the apertures thereby forming papillae, a common feature in Rubiaceae. In Rubiaceae species amoeboid as well as secretory tapeta are reported. In T. gracilipes it is shown that the tapetum cells possess in all developmental stages characteristics of the secretory type. During microsporogenesis the tapetum cytoplasm undergoes considerable changes which may indicate cycles of hyperactivity. Sporopollenin deposition on the pre‐orbicules is mediated by white lines showing correlations with endexine, annulus and columellae ontogeny. These findings corroborate the idea that orbicule wall development can represent a model to study sporopollenin deposition. At anther dehiscence Ca‐oxalate crystals are released out of the ruptured septum cells into the locule, providing a possible visual signal for pollinators.     

Congeneric phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata: Liolaemini)

In poorly known groups for which data are insufficient to develop biologically plausible model‐based approaches to phylogeographical analyses, a ‘first hypotheses’ protocol is suggested as offering the best way to generate hypotheses for subsequent model‐based tests. Preliminary hypotheses are formulated about species boundaries and population processes in three species complexes of the Liolaemus boulengeri group in the context of mtDNA ‘congeneric phylogeography’. The temperate South American Liolaemus provides a model with ancient and recent allopatric divergence across ecologically and geologically complex landscapes, incipient speciation, secondary contact, and discordance between molecular and morphological patterns of variation. Moderately dense sampling of widely distributed ‘inertial’ species in the Patagonian Steppe has revealed hidden genetic and probably species diversity, and also hinted at demographic and historical processes that may have shaped the histories of these taxa. Five of the seven focal species of the present study were paraphyletic for mtDNA genealogies, suggesting that they represent complexes of species, and nested clade phylogeographical analysis (NCPA) analyses suggest that different historical and demographic processes have shaped the observed patterns. Introgression and incomplete lineage sorting are hypothesized as being the cause of some of the observed paraphyly. Provisional delimitations of species are proposed and NCPA is used to generate hypotheses of population history, all of which are subject to further testing. Multi‐faceted studies, involving phylogenetic assessments of independent molecular markers and morphological variation across codistributed taxa with estimates of niche breadths in a landscape context, will likely yield the most promising returns for cross‐validation of hypotheses of population and speciation histories. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 241–275.     

Patterns of asexual reproduction in the fissiparous seastar Coscinasterias acutispina (Asteroidea: Echinodermata) in Japan

The morphological and anatomical characteristics of the fissiparous seastar Coscinasterias acutispina were investigated at 23 sites around Japan in order to clarify the environmental factors determining asexual reproduction. The frequency of recent fission among the sites differed, although no significant differences were evident among the three major sea areas around Japan (the Sea of Japan, Seto Inland Sea, and Pacific Ocean), indicating that the spatial variability in incidence of fission might be determined on a local scale. Morphological observations revealed that the seastars endoparasitized by Dendrogaster okadai had not undergone fission recently, suggesting that fission was suppressed by parasitism. Stepwise multiple regression analysis of the frequency of seastars that had recently undergone fission revealed that nutrient conditions affected the incidence of fission. The relationships between the pyloric caeca index and arm length ratio at the individual level suggested that good nutritional conditions were required before the occurrence of fission. Additionally, regression analysis showed that high temperature was one of the triggers of fission in C. acutispina. Thus, the process of asexual reproduction in C. acutispina appears to be governed by intricate interactions among several factors, including parasitism by D. okadai, nutrient conditions, and high temperature.