Embryo retention,character optimization,and the origin of the extra‐embryonic membranes of the amniotic egg

This study summarizes the data relevant to understanding the appearance of the amniotic egg and provides new analyses to determine the inferences that can be drawn from these data. A survey of the recent literature shows that a consensus exists on the probably primitive absence of extended embryo retention in caecilians, despite recent suggestions to the contrary. The two most recent studies on the evolution of embryo retention in sarcopterygians both suggest that early amniotes lacked extended embryo retention. New analyses of length distribution of random trees suggest that the data on embryo retention do not include a phylogenetic signal, and this implies that character optimization of these data does not yield reliable information on the primitive condition for amniotes. Thus, the study of ancestral features of amniotes will probably have to exploit data from the fossil record.     

Extended embryo retention,caecilian oviparity and amniote origins

A recent discussion of the evolution of the amniotic egg (Laurin and Reisz, 1997) was criticized by Wilkinson and Nussbaum (1998), and these criticisms provoked a rebuttal (Laurin et al., 2000). Here we show that the objections raised by Laurin et al. (2000) do not substantiate the conclusions of Laurin and Reisz (1997). Wealso discuss additional evidence on the ancestral ontogeny of caecilians from the literature. This evidence is inconsistent with the view that extended embryo retention is the ancestral condition for caecilians and that it is a parsimonious interpretation of the condition of the ancestral amniote as argued by Laurin and Reisz (1997) and by Laurin et al. (2000). The available data are more consistent with the traditional hypothesis that the amniotic egg originated as an adaptation of eggs to the terrestrial environment. We also discuss problems in the definition of ontogenetic characters reflecting variation with respect to extended embryo retention, and we present new observations on the early development of the caecilian Gegeneophis ramaswamii Taylor.     

Caecilian viviparity and amniote origins

A recent evaluation of alternative hypotheses for the origin of the amniotic egg, by mapping a single reproductive-mode character onto a phylogeny of tetrapods, concluded that the alternative hypotheses were equally parsimonious. However, this interpretation is dependent upon a mistaken coding of the caecilian amphibians as showing extended embryo retention. Although some caecilians are viviparous, phylogenetic analyses indicate that oviparity is ancestral for the group. With the coding of caecilians corrected, the most parsimonious inference is that the ancestral amniotes did not practice extended embryo retention. A review of the available data indicates that the widespread view that a majority of caecilians are viviparous is mistaken. Oviparity is the dominant reproductive mode in caecilians as it is in other living amphibians.     

Life history of Subulina octona (Brugüière) (Gastropoda: Pulmonata: Subulinidae) based on four-year laboratory observations and a comparative histological analysis of egg-retaining and ovoviviparous subulinids

In the present study, we aimed to characterize the life history of the land snail Subulina octona integrating information on life-history traits and morphology. We also compared the histology of the free-oviduct and spermoviduct of ovoviviparous and egg-retaining species of subulinids. We considered as ovoviviparous the species in which the complete embryonic development as well as egg hatching occurs inside the parent’s body and, at the end of this process, the parent releases juveniles instead of eggs. We considered as egg-retaining the species in which a great part of the embryonic development takes place inside the parent’s body and the eggs laid contain well-developed embryos. The results showed that the free-oviduct of both ovoviviparous and egg-retaining species shows a histological arrangement that confers greater strength to its walls and is probably related to egg retention. The wall of the spermoviduct is formed by pseudostratified columnar epithelium, with cell apical processes (probably cilia), and by underlying secretory cells. In gravid egg-retaining snails, the eggshells appear in close contact with the secretory cells of the spermoviduct. This fact suggests that these cells play a role in eggshell formation. The present study is the first account for histological features of S. octona, Allopeas gracile and Allopeas micra. The life history of S. octona is a combination of long lifespan, early sexual maturity, indeterminate growth and egg retention. Egg retention limits the number of young that can be produced in one reproductive event. However, the reproductive strategy adopted by S. octona, associating egg retention and K-strategism, compensates this cost of retaining eggs, because the reproductive success may be enhanced as a result of the higher survival of juveniles and the possibility of performing several reproductive events during the year.     

The evolution of nest building and communal nesting in Ageniellini (Insecta: Hymenoptera: Pompilidae)

Pompilid wasps prey upon spiders and use a single spider per nest cell. The majority of species dig simple burrows in the soil in which to cache their spider, on which an egg is laid. The nest is not revisited, but another nest is prepared elsewhere. However, members of the tribe Ageniellini show much diversity in nesting behaviour. Species of the genus Ageniella nest in pre-existing cavities in the soil, closing off their cells with bits of debris. Most other Ageniellini that have been studied carry water to make mud pellets, from which ovoid nest cells are made. Usually a series of such cells is made in close proximity, often under loose bark or stones or in hollow stems. A few species make free nests above ground, usually in protected places or with thick mud walls. In some cases nest sites are known to be re-occupied by members of successive generations. Several species are known to nest communally, co-operating in nest defense and in cell building. In many ways the evolution of nesting behaviour in this group parallels that in the mud-using Vespidae, but the use of a single prey per cell precludes development of progressive provisioning and of eusociality similar to that of many Vespidae.     

Comparative investigation of the late embryogenesis of Leptodora kindtii (Focke, 1844) (Crustacea: Branchiopoda), with notes on types of embryonic development and larvae in Cladocera

The embryonic development of the parthenogenetic eggs of Leptodora kindtii has been investigated by observing living embryos removed from the female brood pouch. The sequence of morphological changes was analysed, as was the time at which the activity of certain organs began. The timing of these events at 20–22°C is documented. These data were compared with similar information obtained for some representatives of Ctenopoda and Anomopoda. The sequence of appearance of rudiments of organs on the ventral side of the embryos is similar in all these groups but the timing of shedding of the embryonic membranes differs. Hatching from the inner egg membranes takes place at the late metanauplius stage in Leptodora as in Ctenopoda and Cyclestherida, while in Anomopoda this stage is embryonized. The development of Leptodora is more like that of Ctenopoda than of Anomopoda. The embryos of all the above‐mentioned groups begin to grow at approximately the same morphological stage. A new scheme for the periodization of the embryogenesis of L. kindtii is proposed, which includes four well‐separated stages. Previously, the same number of developmental stages has been reported for Ctenopoda and Anomopoda. In Leptodora, as well as in Ctenopoda, the first two of these occur within the egg membranes. Cyclestheria has at least one embryonic moult and one embryonic stage more than Leptodora and other cladocerans. Embryonic stages 3 and 4 of the parthenogenetic eggs of L. kindtii represent a peculiar type of secondary larvae developing inside the female's brood pouch. The appearance of a free‐living nauplius in L. kindtii arises from its mode of hatching from the gamogenetic egg.     

On the natural history of the South American pepper frog,Leptodactylus labyrinthicus (Spix, 1824) (Anura: Leptodactylidae)

Some species of Leptodactylus of the L. pentadactylus group lay their eggs outside water but the tadpoles need to reach water to complete the larval phase; other species complete development in terrestrial nests. Here we present details of the reproduction of L. labyrinthicus in south‐eastern Brazil. The proportion of tadpoles and trophic eggs in aged egg clutches was determined, as well as the growth of the tadpoles while in the nest. The gut contents of tadpoles that were in egg clutches of frogs were analysed. Adult males did not differ from females in size and had hypertrophied forearms and an enlarged spine on the thumb. Reproduction was initiated with the first rains of August/September and extended to mid‐January. Calling and spawning occurred at permanent or temporary water bodies. The foam nests were built in excavated basins outside of, but close to the water. The male determined the place of the basin construction; after amplexus, the female completed the excavation. The amplexus was axillary. One female spent the day after spawning in the foam. The eggs were pale grey, the yolk averaging 2.3?mm in diameter. The mean number of eggs was 2101 per egg clutch. The number of tadpoles in individual nests varied between 0.05% and 11.40% in relation to the total laid eggs. The tadpoles entered water when rains flooded the basin. The tadpoles grew to 12 times the weight of an individual egg while in the nest; no nesting tadpole was beyond stage 25. The longest time we followed tadpoles in a nest was 25 days. Tadpoles were found preying upon eggs of three other frog species and upon conspecific eggs. Males fought by grasping each other in a belly‐to‐belly position; the powerful arms and the thumb spines represent weapons. Even though males can reach maturity in the season following birth, small size would prevent them from establishing their own territory. All the species of the L. pentadactylus group may build their foam nests within excavated basins. The basins may protect the eggs and embryos from cannibalistic tadpoles and may have an anti‐desiccation effect. In order to produce trophic eggs, the female may delay laying additional unfertilized eggs until after the male has abandoned the foam nest. Anuran eggs represent an important food item for tadpoles after they leave the nest.     

Spawning behaviour of Engystomops pustulatus (Anura: Leptodactylidae)

Foam nests have evolved independently in several amphibian groups as an adaptive response to prevent predation and desiccation in dry environments. Nests are normally laid on ponds, or in underground galleries, humid forest leaf litter or terrestrial bromeliads. They are built when males or females beat a foam precursor associated with the egg masses extruded by the female. The spawning process requires the synchronic actions of the mating pair to obtain a hemispheric nest that protects the offspring. Herein, we describe the spawning behaviour of Engystomops pustulatus based on videos from 13 nesting couples from the lowlands of western Ecuador. Three variables were measured as indicators of male effort: duration of mixing events, duration of resting periods, and number of kicks per mixing event. We consider that not only male physical effort but also female behaviour influences nest structure. We suggest that nest building requires prolonged and intense physical activity by the male as well as the female’s steady position during spawning and female’s oviposition site selection. Nest building has two phases. In the first phase, the duration of resting periods, the duration of mixing events, and the number of kicks increase and are highly variable. During the second phase the three variables stabilise until the end. The volume of the nest increased mainly during the second phase. In four nesting events we observed kicking movements by the female. To our knowledge, this is the first time that female kicking has been observed in leptodactylid frogs. The function of this behaviour is unknown but our observations suggest that it may be triggered by insufficient male effort. Traditionally, female mate choice in Engystomops has been explained under models of indirect benefits exclusively. We argue that the prolonged male activity during nesting could influence female fitness directly. This will allow the operation of sexual selection via direct benefits.     

A classification and phylogeny of post-Palaeozoic sea stars (Asteroidea: Echinodermata)

A revised classification and phylogeny at the family level and above are presented for post-Palaeozoic sea stars. Monophyly of the group is established by a character suite taken from the ambulacral column that thus far has been recognized in only one Palaeozoic genus. Compared to earlier studies, character selection here placed relatively greater emphasis on morphology and arrangement of ossicles and ossicular systems. Functional implications of many features are surveyed. Thirty-four families, three extinct, are recognized and a number of older familial concepts are suppressed; the extinct Trichasteropsidae is proposed. Superfamilies are recognized for the Valvatida. Eight orders, including the new, extinct Trichasteropsida, and three superorders are recognized. No living sea star is primitive in the sense of being close to ancestral sea stars and other echinoderm groups; the Paxillosida, which commonly has been considered primitive, is here considered specialized.     

Reproductive characteristics of Elaphe quadrivirgata (Serpentes: Colubridae) from ecologically dissimilar main island and island populations

We compared reproductive characteristics of a snake (Elaphe quadrivirgata) from two geographically distinct populations, which differ considerably in habitat use and diet. Snakes in Shiga Prefecture, in the central western area of the main island of Japan, occupy open rice fields and predominantly eat frogs. Snakes on Yakushima Island, in southwestern Japan, occupy a forest environment and mainly eat lizards. Maternal body size of snakes from Yakushima was significantly smaller than that of snakes from Shiga. Oviposition date was significantly later in snakes from Yakushima than in those from Shiga, and the former laid a smaller number of considerably elongated eggs than the latter. Relative clutch mass was significantly higher in the Shiga than in the Yakushima population, but egg mass in relation to maternal body mass of the latter was larger than that of the former. Hatchlings of Yakushima were larger than those of Shiga. This size difference in hatchlings was not fully explained by differences in egg size. These differences in reproductive characteristics between the two populations do not seem to be a consequence of morphological constraints of dam snakes. Instead, we presume that the differences in reproductive characteristics reflect local adaptations.     

Activity and reproduction in Megalobulimus paranaguensis (Gastropoda,Eupulmonata): implications for conservation in captivity for a South American land snail

ABSTRACT

In South America, Megalobulimus includes a number of threatened species, as well the largest land snails on the continent. The activity patterns and reproductive aspects of this group have not been documented. This work describes the daily and seasonal activity patterns and reproduction of M. paranaguensis. We maintained specimens in the laboratory for one year, and we quantified their behaviour for one hour at four different times of the day (0 h, 6 h, 12 h and 18 h) during three days in four months (August, September, April and May). The number of postures, hatching rate, time of hatching since oviposition and mortality rate among juveniles for each month were also quantified. Megalobulimus paranaguensis was more active in August, and had a egg laying peak one month after. Fifty-one eggs were laid by 12 captive individuals throughout the year, with a mean value of 4.25 eggs per individual. The hatching rate was 80.39%, and the time of hatching since oviposition was 56.7 ± 4.3 days. In two eggs, we observed the presence of twins. The mortality rate among juveniles was low (9.30%) indicating that rearing land snails in captivity has the potential to be an important and viable tool for the management and conservation of these organisms.     

Reproductive biology and strategies of nine meloid beetles from Central Europe (Coleoptera: Meloidae)

The reproductive biology of meloid species from Central Europe is investigated by means of laboratory breeding experiments. They show that the total reproductive potential of meloids, with up to 40,000 eggs, largely exceeds prior data. Furthermore, the number of laid eggs strongly relates to the way by which the triungulins find their host. Clutch size correlates significantly with the size of the beetle, while egg size is independent of this. Both clutch size and egg size decrease with each oviposition. For some species, reproductive data are used to demonstrate how they have adapted to their habitat and to point out existing trade‐offs. Based on the obtained results and on an evaluation of the relevant literature, three types of reproductive strategy can be distinguished within the meloids: (1) very high reproductive rates of open field species with phoretic larvae; (2) average reproductive rates of forest populations of species with phoretic larvae and of species the larvae of which search the nests of their hosts actively; and (3) small to average reproductive rates of those species depositing their clutches in the immediate proximity of their hosts' nests.     

On the life-habits and developmental stages of Nidomyia cana Papp (Diptera,Borboropsidae)

Eggs, second and third instar larvae of the enigmatic species Nidomyia cana, which was discovered most recently from the nests of raptorial birds in southern Hungary, are described. Both egg and larvae show peculiarities among the so-called heleomyzoid flies. Larvae were found in wet vegetable material (mostly pieces of poplar bark) of a buzzard nest and their shape is of a saprophagous type. Females are macro-oviparous, with a maximum of 15 to 17 eggs laid in one batch. Adults live mostly on the surface the of the nest, they copulate there, and they find shelter in the nest itself. No direct contact with adults or nestlings within the nest was observed. It is suggested that overwintering occurs as pupae (pharate adults) underground, beneath the nest. However, if N. cana lives exclusively in the nests of raptors, adults would have difficulties in finding a new nest, and it is probable that they are transferred to new nests attached to the birds.     

New insights on the external features of egg capsules and embryo development in the squid Loligo vulgaris

The embryonic development of the squid Loligo vulgaris was observed from 183 egg masses collected from special devices deployed throughout Cabrera National Park (Baleares Islands, western Mediterranean Sea). Sequence alignment analysis of the cytochrome oxidase I gene revealed that all embryos belonged to L. vulgaris. In total, 549 egg capsules were examined. Viable egg capsules (n = 420) were classified into one of five maturation stages according to the primary external features. The length of the viable egg capsules varied between 40 and 170 mm, and increased with embryonic development. The non-viable capsules (n = 129) were categorized into four groups: I (Ginger root), non-viable II and III, and empty egg capsule (IV). The percentage of non-viable capsules (i.e. grades I, II and III) was 92.25%. Empty capsules accounted for 7.75% of the total non-viable egg capsules. Embryonic development was classified into a second scale of eight stages. Egg capsule stage and embryonic stage were significantly related (n = 420; p < 0.001), facilitating the determination of the embryo developmental phase based on the outward appearance of the egg capsules. The embryo development stage based on the external features of the egg capsules might constitute an innovative tool for in situ embryological data collection. This new method is neither time consuming nor invasive, and could be helpful in fishing cruises, for scuba diving visual census in natural habitats and for laboratory culture. Slight variability in the developmental embryonic stages within egg capsules from the same egg mass was identified. The origin of this asynchrony is discussed. Chronological appearance of organs was similar to that of the six loliginid species previously examined. However, some developmental changes in the timing or rate of events (heterochronies) were observed: Hoyle’s organ was formed earlier in L. vulgaris and the appearance of ventral chromatophores was slightly delayed (2 days) compared with the other species considered.     

XCII.—The early lophophore and subsequent loop (Brachiopoda)

Abstract

Examination of recent brachiopods, Terebratulina, Terebratella and Eucalathis, illustrates earlier observations that the young lophophores are centrifugal in Terebratulacea and centripetal in Terebratellacea, and suggests that both median septa and certain median structures free of the valve-floor are initiated by centripetal lophophores. This is turn suggests that the absence of the centronella-stage in Cancellothyridae, etc., is due not to tachygenesis but to different early lophophores from those of the Centronellidae, etc.; that the abnormal loops of Hamptonina may have been due to abnormal lophophore-orientation as well as retarded development; and that lophophore mutation could occasion changes of up to superfamily rank without leaving intermediate forms. It is concluded from consideration of the loop-lophophore relationship discussed above, against the background of present knowledge of Palaeozoic and certain Triassic loop-bearing brachiopods, that such earlier forms are neither Terebratulacea (in which they are usually placed) nor Terebratellacea, but that after re-study it should be possible to group them as superfamilies in which the brachial structures have less value for classification than in the Mesozoic superfamilies quoted.     

Development and demography of Phasmahyla jandaia (Bokermann and Sazima, 1978) (Anura,Hylidae) tadpoles in an Atlantic Forest site,southeastern Brazil

Mortality rates are often high during the anuran tadpole stage, making demographic parameters in this phase of their life cycle important to the understanding of population demography. Nevertheless, information on mortality rates or population size is lacking for most Brazilian anurans. Herein we report on the demographic parameters of Phasmahyla jandaia (Hylidae) during the larval stage, including number of individuals through time, mortality rates and time of recruitment. Tadpole recruitment occurred year‐round, although froglet recruitment to the terrestrial habitat was restricted to the warm, rainy season. This trend seems to result in high plasticity of the tadpole developmental period. Mortality rates decreased with advancing tadpole stage, totalling 77% during the whole aquatic phase. Survivorship can be regarded as intermediate compared with other anurans for which data are available, and is in accordance with a high investment in egg size and the low number of eggs laid by P. jandaia.     

Postembryonic development and population parameters of Alpaida veniliae (Araneae,Araneidae), reared in the laboratory

We studied the postembryonic development from hatching to the adult stage and determined population parameters of Alpaida veniliae. It is one of the most abundant species of the orb-weaving guild of the spider community of soybean crops in Buenos Aires province, Argentina. The first three instars occurred inside the egg sac, and instar IV (spiderlings) started the dispersion from it. The female of Alpaida veniliae achieved greater adult size and adult longevity than the male. Mean fecundity, mean number of egg sacs per female, mean number of eggs per egg sac, as well as the net reproductive rate, the intrinsic rate of increase (r), the generation time (T), and the reproductive values (V _x) of three cohorts were determined under laboratory conditions, indicating a high capacity for growth. Their biological and ecological attributes indicate the importance of conservation of this predator as a natural enemy of soybean crop pests.     

Fossil evidence for the escalation and origin of marine mutualisms

Mutualism is an interactive, mutually beneficial relationship between two or rarely more unrelated taxa. It is ubiquitous in most environments. In the marine environment it is most likely to occur in shallow marine, oligotrophic regions, the open marine photic zone and oxygen-limited environments. Whereas the eukaryote cell is of some antiquity, other mutualisms may have been acquired during the Phanerozoic. The fossil record provides inferential evidence for the minimum time of origin and occurrence of mutualisms. This suggests that a variety of mutualisms may have occurred during the Palaeozoic with most apparently being terminated by the Permian mass extinction. Tracing back the origin of extant mutualisms suggests that the majority of these began in the Mesozoic or Cenozoic and indicate an increase to the present day. Whether the increase in mutualisms is a product of increased biodiversity or whether mutualisms may have partly driven biodiversity is unclear.