The reproductive biology of some serolid isopods from the Antarctic

Summary Several species of serolid isopod from Signy Island, South Orkney Islands and also from South Georgia were studied, and aspects of their reproductive behaviour and associated morphology are discussed. InSerolis polita Pfeffer egg-laying took place at a mean age of 28 months, and the eggs were then incubated in a ventral marsupium for a period of 20 months before the release of juveniles in the spring. The mean egg dry weight at spawning was 0.68 mg forS. polita and 3.35 mg forSerolis cornuta Studer. The egg weight increased during marsupial development due to the uptake of minerals, but the total calorific content fell. Within each species brood size was linearly related to female size and the mean number of eggs varied from 40 forS. polita to 232 forSerolis pagenstecheri Pfeffer. There was no detectable marsupial mortality.S. polita appeared to breed repeatedly at two-year intervals, butS. cornuta probably only breeds once. The reproductive biology of Antarctic serolids is compared with that of temperate isopods, and it is shown that they produce larger young which is related to their larger adult size. The extremely protracted period of marsupial incubation is associated with the need to synchronize reproduction with the seasonal nature of Antarctic primary production.     

Do Seaducks Minimise the Flightless Period?: Inter- and Intra-Specific Comparisons of Remigial Moult

Remigial moult is one of the crucial events in the annual life cycle of waterfowl as it is energetically costly, lasts several weeks, and is a period of high vulnerability due to flightlessness. In waterfowl, remigial moult can be considered as an energy-predation trade-off, meaning that heavier individuals would minimise the flightless period by increasing feather growth rate and energy expenditure. Alternatively, they could reduce body mass at the end of this period, thereby reducing wing-loading to increase flight capability. We studied timing of remigial moult, primary growth rates, flightlessness duration, and the pattern of body mass variation in 5 species of captive seaducks (Melanitta fusca, M. perspicillata, Clangula hyemalis, Histrionicus histrionicus, and Somateria mollissima) ranging in size from 0.5 to 2.0 kg. Their feather growth rates weakly increased with body mass (M^0.059) and no correlation was found at the intra-specific level. Consequently, heavier seaduck species and especially heavier individuals had a longer flightless period. Although birds had access to food ad libidum, body mass first increased then decreased, the latter coinciding with maximum feather growth rate. Level of body mass when birds regained flight ability was similar to level observed at the beginning of remigial moult, suggesting they were not using a strategic reduction of body mass to reduce the flightlessness duration. We suggest that the moulting strategy of seaducks may be the result of a compromise between using an intense moult strategy (simultaneous moult) and a low feather growth rate without prejudice to feather quality. Despite the controlled captive status of the studied seaducks, all five species as well as both sexes within each species showed timing of moult reflecting that of wild birds, suggesting there is a genetic component acting to shape moult timing within wild birds.     

Moult cycle and growth of the crab Halicarcinus planatus (Brachyura, Hymenosomatidae) in the Beagle Channel, southern tip of South America

The crab Halicarcinus planatus is the only hymenosomatid crab that inhabits the southern tip of South America and is the only decapod species that reproduces twice a year in the Beagle Channel. In this article, we study the moult cycle in the field (moult frequency, analysis of size frequency distribution) and linked it with growth studied in the laboratory (absolute and per cent growth increment, Hiatt function). Hiatt functions were similar for males and females. Moult frequency was seasonal: in early austral spring and in austral summer. In females, the pubertal moult is the terminal moult, whereas males continue moulting after attaining the size of morphometric maturity. Moult increment was highly variable. The relationship between absolute moult increment and crab size was described by a quadratic function. Per cent growth increment decreased with size, and relationships were different for each sex: linear for females and quadratic for males. Seven and eight modal groups explained the size frequency distributions for females and males from the field, respectively, and revealed the existence of two cohorts of recruits per year. Further modal analysis was mainly hampered by the high variability of size increment that could make any moulting individual fall in its own or one of two following modal groups. The antagonism between growth and reproduction was evident in small males. We hypothesize that the terminal pubertal moult is an advantageous feature that allows females to maximize their investment in reproduction after their terminal moult, which allows this species to have two spawnings per year.     

Wood anatomy and growth rate of seasonally dry tropical forest trees in the Marañón River Valley,northern Peru

Seasonally dry tropical forests (SDTF) currently occupy only 10% of the original area of their natural distribution. In the Marañón river valley, north western of Peru, occurs one of the most important SDTF of the neotropics. It has an endemic woody flora that represent 32% of the species, being threatened by its increasing fragmentation, and the knowledge of the ecology, forestry and conservation of the tree species is scarce. In this context this work presents the result of evaluation of the wood anatomy with emphasis to growth ring analysis, dating trees and stem growth rate determination of four tree species: Anadenanthera colubrina (Vell.) Brenan, Cedrela kuelapensis T.D. Penn. & Daza, Cordia iguaguana Melch. Ex I.M. Johnst., and Esenbeckia cornuta Eng., the last three being endemic and threatened with extinction. Our results determined that Cedrela kuelapensis showed the highest radial growth of the stem in relation to the other congeners of the neotropical region. Lower stem growth was observed for E. cornuta trees, according to the pattern of other tropical species. The results also indicate that populations of C. iguaguana and Esenbeckia cornuta in the zone are mainly composed by young trees exhibiting strong anthropic pressure due to the quality of their timber and, in consequence, more efforts are needed in the management of these resources to guarantee their sustainability.     

In vitro culture of hymenopterous larval endoparasitoids

The solitary endoparasitoids Cotesia (= Apanteles) marginiventris and Microplitis croceipes, which attack noctuid larvae, were successfully cultured from post-germband stage eggs to fully grown first-instar larvae in completely artificial (but undefined) media. However, growth rates were slower than in vivo, and while some larvae of each species attained apparent competence to moult (based upon characteristic size and form), none did so.Up to 75% of pregermband M. croceipes eggs developed and hatched when co-cultured with fat body from non-parasitized larvae of the habitual host, Heliothis zea. Heat-treated haemolymph plasma from a non-permissive host, Manduca sexta, also fostered development by ca. 40% of newly laid M. croceipes eggs, but plasma from H. zea larvae was not beneficial. Growth rates of C. marginiventris and M. croceipes larvae were not improved by haemolymph plasma from their habitual hosts (Spodoptera frugiperda and H. zea, respectively).Heat-treated foetal bovine serum was growth-promoting for post-germband eggs and larvae of both parasitoids. Treatment of foetal bovine serum by charcoal adsorption (to remove sterols and peptide hormones) diminished its value as a supplement. Two major proteins of the serum, foetuin and albumin, also promoted survival and growth for both parasitoid species. The potential relevance of these findings to in vitro rearing of endoparasitoids is discussed.     

The opercular moult in Spirorbis spirorbis (L.) and S. pusilloides Bush (Polychaeta: Serpulidae)

Opercular moult has been examined in two spirorbids, the tube-incubating Spirorbis spirorbis (L.) and the opercular incubating Spirorbis pusilloides Bush. In both species epithelial ingrowth cuts off the old operculum and a new one differentiates on the proximal side of the constriction. The immediate fate of the severed tissues differs in the two species; in S. spirorbis the epithelia slowly wither as the platelike operculum finally breaks away by rupture of the cuticle while in S. pusilloides there is a cataclysmic autolysis of the epithelial tissues although the cuticle is retained and functions as an incubation chamber which ruptures at the close of incubation to release the larvae and is then shortly discarded.Opercular moult, rare in the immature animal in either species, normally begins with the onset of breeding. In S. pusilloides it is an essential preliminary to brooding and throughout the breeding season opercular moult rigidly alternates with brood incubation to produce a new incubation chamber for each successive brood. In S. spirorbis there is also some evidence of a partial correlation between brooding and opercular moult but the two events overlap and moulting is less frequent than the production of broods. A brief consideration is given to the evolution of the brooding habit and the tentative conclusion reached that opercular incubation has evolved twice in the Spirorbinae.     

Cryptic introgression into the kidney saxifrage (Saxifraga hirsuta) from its more abundant sympatric congener Saxifraga spathularis,and the potential risk of genetic assimilation

Background and Aims Although hybridization can play a positive role in plant evolution, it has been shown that excessive unidirectional hybridization can result in replacement of a species’ gene pool, and even the extinction of rare species via genetic assimilation. This study examines levels of introgression between the common Saxifraga spathularis and its rarer congener S. hirsuta, which have been observed to hybridize in the wild where they occur sympatrically.Methods Seven species-specific single nucleotide polymorphisms (SNPs) were analysed in 1025 plants representing both species and their hybrid, S. × polita, from 29 sites across their ranges in Ireland. In addition, species distribution modelling was carried out to determine whether the relative abundance of the two parental species is likely to change under future climate scenarios.Key Results Saxifraga spathularis individuals tended to be genetically pure, exhibiting little or no introgression from S. hirsuta, but significant levels of introgression of S. spathularis alleles into S. hirsuta were observed, indicating that populations exhibiting S. hirsuta morphology are more like a hybrid swarm, consisting of backcrosses and F₂s. Populations of the hybrid, S. × polita, were generally comprised of F₁s or F₂s, with some evidence of backcrossing. Species distribution modelling under projected future climate scenarios indicated an increase in suitable habitats for both parental species.Conclusions Levels of introgression observed in this study in both S. spathularis and S. hirsuta would appear to be correlated with the relative abundance of the species. Significant introgression of S. spathularis alleles was detected in the majority of the S. hirsuta populations analysed and, consequently, ongoing introgression would appear to represent a threat to the genetic integrity of S. hirsuta, particularly in areas where the species exists sympatrically with its congener and where it is greatly outnumbered.     

Molecular Evidence for the Hybrid Origin of Ilex dabieshanensis (Aquifoliaceae)

Ilex, the largest genus of dioecious woody plants, is a good study system to assess the role of hybridization in speciation and evolution. Ilex dabieshanensis, a tree endemic to Dabieshan Mountains region, was initially described as a new species. Based on morphological intermediacy and sympatric distribution with its putative parental species, I. cornuta and I. latifolia, we proposed it as a natural hybrid between them. In this study, we sequenced one chloroplast intergenic spacer (trnH-psbA) and two nuclear genes (gapC and nepGS) in I. dabieshanensis and its putative parental species to test the hybrid origin hypothesis. Our results showed that there were one to two differentially fixed sequence differences between I. cornuta and I. latifolia at the two nuclear genes. Twelve of the 14 individuals of I. dabieshanensis exhibited additivity in chromatograms on these differentially fixed sites at both nuclear genes, and the remaining two exhibited additivity in chromatograms on the fixed site at only the nepGS gene. Except one haplotype of I. cornuta at the nepGS gene, all of the haplotypes of I. cornuta at the two nuclear genes were well separated from those of I. latifolia, and most haplotypes of I. dabieshanensis were shared with those of I. cornuta and I. latifolia. Phylogenetic analysis of these haplotypes was largely consistent with haplotype network analysis. I. cornuta and I. latifolia differed by two nucleotide substitutions in the chloroplast intergenic spacer, and 12 individuals of I. dabieshanensis had the same sequences as I. latifolia, while the remaining two were identical with I. cornuta. The molecular data provide convincing evidence for the hybrid origin of I. dabieshanensis and asymmetrical direction of hybridization. One haplotype of I. cornuta at the nepGS gene was nested with those of I. latifolia, indicating introgression to I. cornuta.     

Growth rate as a measure of food value in thaidid gastropods: Assumptions and implications for prey morphology and distribution

Rates of body growth were measured for three species of rocky intertidal gastropods: Thais (or Nucella) lamellosa (Gmelin), T. (or N.) canaliculata (Duclos), and T. (or N.) emarginata (Deshayes). Three size classes of each predator species were grown experimentally in cages at two tidal heights on four size classes of each prey species: Semibalanus cariosus (Pallas), Balanus glandula Darwin, and Mytilus edulis L. Growth was also assessed for Thais canaliculata feeding on Mytilus californianus Conrad, and Thais emarginata feeding on Chthamalus dalli Pilsbry.Rates of body growth varied as a function of predator size, prey size and prey species. With one exception (large Thais canaliculata) intermediate sized Balanus glandula promoted the most rapid growth for all sizes of all three species of Thais; thus these potentially competing predator species have the same highest ranked prey. Among prey promoting slower growth, those encountered commonly in the normal habitats of the predator promoted more rapid growth than those encountered rarely, suggesting that past evolutionary experience has influenced present food value (= growth potential) of prey. For some predators, the ranking of prey species changed with predator size; Chthamalus dalli and similar sized Balanus glandula promoted comparable growth in small Thais emarginata but Chthamalus dalli promoted much slower growth than similar sized Balanus glandula in larger Thais emarginata. Rank differences based on prey size in Balanusglandula became more pronounced and in some instances changed with increasing predator size; larger B. glandula promoted relatively faster growth for large snails than for small snails in all Thais species. Finally, growth rates were correlated with two important attributes of fitness in T. canaliculata and T. emarginata; in general, prey promoting more rapid growth also resulted in an earlier age of first reproduction for initially immature snails and higher rate of egg capsule production for mature individuals. Thus, these growth rates provide a basis from which to examine quantitatively patterns of prey selection from an energy- or growth-maximization perspective.These patterns of predator growth also permit inferences about the possible influence of predation on the evolution of prey morphology, life-history, and microhabitat distribution. Size refuges from Thais predation were confirmed for both Semibalanus cariosus and Mytilus californianus. Both Balanus glandula and Mytilus edulis, on the other hand, were vulnerable to predation by all sizes of Thais examined (> 10 mm) regardless of prey size; thus, neither of these species achieves a size refuge. In addition, on rocky shores, vertical distributions of the prey species reflect general preference patterns of their predators: higher value prey species, i.e. those with less well developed defensive morphologies (Balanus glandula, Mytilus edulis), generally occur higher on the shore. This pattern would be expected if preferred prey are consumed first lower on the shore where they are more available.     

Lack of seasonal and moult-related stress modulation in an opportunistically breeding bird: The white-plumed honeyeater (Lichenostomus penicillatus)

This article is part of a Special Issue “SBN 2014”.In most vertebrate species, glucocorticoid levels and stress sensitivity vary in relation to season and life-history stage. In birds, baseline corticosterone (CORT) and stress sensitivity are typically highest while breeding and decrease substantially during moult. Because elevated CORT adversely affects protein synthesis, moult-related CORT suppression is thought to be necessary for forming high-quality feathers. Surprisingly, some passerine species lack moult-related CORT suppression, but these are distinguished by having slow rates of moult and being opportunistic breeders. We examined baseline and stress-induced CORT levels in an opportunistically breeding Australian passerine, the white-plumed honeyeater (Lichenostomus penicillatus). Although this species has a slower moult rate than high-latitude breeders, it differs little from north-temperate passerines. Neither baseline nor stress-induced CORT levels varied with season (winter, spring or summer), sex or moult status in adult birds. While breeding tended to be highest in early spring through late summer, laparotomies revealed only limited reduction in testicular size in males the year round. In all but one sampling period, at least some females displayed follicular hierarchy. Breeding usually coincides with outbreaks of phytophagous insects, which can happen at any time of the year. This results in moult/breeding overlap when infestations occur in late spring or summer. The ability of this species to moult and breed at the same time while having breeding-levels of CORT demonstrates that CORT suppression is not a prerequisite for synthesis of high-quality feathers. An experimental design incorporating moulting and non-moulting phenotypes is suggested to test the functional significance of CORT suppression in other species.     

Do closely related species share of feeding niche along growth? Diets of three sympatric species of the mojarras (Actinopterygii: Gerreidae) in a tropical bay in southeastern Brazil

Understanding the trophic relationships among closely related species is a way to obtain subsidies for their management and conservation of their habitats. The diets of three co-occurring abundant fish species of the Gerreidae family (Diapterus rhombeus, Eucinostomus argenteus and Eucinostomus gula) in a tropical bay were described. The tested hypothesis was that the three sympatric species present shifts in their use of resource during the ontogenetic development to facilitate their coexistence. Size groups for each species were categorized according to breakpoints in the morphological structures determined by piecewise regression models. Significant overlapping in diets was found for all size classes of D. rhombeus but not for size classes of the Eucinostomus genus. Furthermore, different size classes of D. rhombeus did not overlap diet with size classes of the Eucinostomus genus. The specialization in feeding niches corresponding to growth seems to bring benefits for this group of fish rather than a generalist feeding strategy. The hypothesis of the available resources partitioning was accepted only between the two genera (Diapterus and Eucinostomus), and among size classes of the Eucinostomus genus that seemed to follow the principle of limiting similarity. However, different size classes of D. rhombeus exhibited strong evidence of an intraspecific overlapping of the trophic niche. It seems that different processes related to use of the trophic niche dimension are structuring these closely related fish species.     

Moult cycle and its chronology in Mysis mixta and Neomysis integer (Crustacea, Mysidacea): implications for growth assessment

The moult-staging technique was used to determine the main moult stages (MS) and their duration in laboratory-reared juveniles and subadults of Mysis mixta and Neomysis integer. The relative duration of each stage was similar for both species with premoult occupying the major part of the moult cycle (51% and 44% for M. mixta and N. integer, respectively), followed by postmoult (26% and 34%) and intermoult (23% and 22%). Effects of temperature and feeding regimes on the chronology of the moult cycle were investigated. When the duration of the moult cycle (MCD) was extended by manipulating the feeding regime, animals prolonged their late postmoult and early premoult stages. At 12 and 5 °C, no specific moult stage varied in relative duration as long as food supply was high. Field application of moult staging for growth assessment was tested using wild-caught M. mixta. The MCD estimated via moulting experiments was compared to that obtained by analyzing moult stage distribution combined with the experimentally obtained data on stage duration. Close correspondence between two methods of MCD assessment was observed with moult-staging technique being particularly useful in situations when conducting of experiments immediately after collection is not feasible.     

Postembryonic growth in two species of freshwater Ostracoda (Crustacea) shows a size-age sigmoid model fit and temperature effects on development time, but no clear temperature-size rule (TSR) pattern

Although the temperature-size rule is a widespread phenomenon that describes the impact of temperature on the intraspecific size of ectotherms, what determines the wide intrinsic variation in moult increment and the intermoult period within and between Crustacea species remains unknown. This work characterizes the growth of freshwater ostracods Chlamydotheca incisa and Strandesia bicuspis under different controlled temperatures and identifies growth patterns. Animals were collected from temporary ponds in Argentina. The experiment was done at two constant temperatures: 17 and 23 °C. The intermoult time and the time from hatching to the final moult were calculated. Three different growth regression equations were tested: von Bertalanffy and two sigmoidal models (Sigmoid and Gompertz). For both species, significant differences in the duration of each instar were found by comparing individuals grown at 17 and at 23 °C. A strong temperature effect was noted on intermoult time but not on growth factors. The best model selected for the size-age relationship was a sigmoid growth type, indicating accelerated growth in the earliest juvenile instars. These results challenge the widely-accepted application of nonsigmoidal growth models and are in agreement with recent analyses of growth patterns in aquatic invertebrates when early juvenile instars are considered.     

Haemolymph constituents and osmolality as functions of moult stage,body weight,and feeding status in marron,Cherax cainii () and yabbies,Cherax destructor (Clark, 1936)

The study investigates the change in osmolality and haemolymph constituents in marron Cherax cainii and yabbies Cherax destructor associated with moult stages, body weights and their feeding status. A total of 582 haemolymph samples from 5 moult stages (postmoult-AB, intermoult-C, and premoult stages – D₀, D₁, D₂), two body weight classes (2–15 g and 61–75 g) and nutritional status were used for analysis of osmolality, protein, glucose, and ionic concentrations of potassium and chloride following the standard biochemical procedures. The haemolymph protein, glucose, potassium and chloride levels were highest at intermoult and early premoult stages, and lowest at postmoult in both crayfish species. Except protein, no significant differences were seen in analyzed parameters between various weight classes and two species. Haemolymph osmolality, protein and glucose were significantly higher in fed crayfish, whereas no variations in haemolymph potassium and chloride concentrations were observed between the fed and unfed crayfish. Maximum osmolality was recorded at 7–8 h after feeding in both crayfish species. The results showed that the biochemical changes in the haemolymph of marron and yabbies are related to moult stages, body weight and feeding and thus can be used as tools for determining suitable diets.     

Experiments on the growth of Penaeus vannamei Boone

Growth rates of Penaeus vannamei Boone were estimated from field and laboratory experiments using both marked and unmarked shrimp. Field results with unmarked shrimp indicated growth rates of between 0.05 mm increase in carapace length per day (mm c.l./day) and 0.17 mm c.l./day, although rates of up to 0.31 mm c.l./day were also recorded. Moult intervals ranged from5.5 to 34 days with a mean of 12.5 days. Moult increment ranged from 0.2 mm c.l./moult to 1.4 mm c.l./moult with an overall mean of 0.7 mm/moult. Growth was negatively correlated with size over all size ranges, with a marked decrease in growth rate occurring at about 28 mm c.l. Shrimp marked with Floy streamer tags generally grew more slowly than unmarked shrimp, but this was exaggerated by the stress of regular handling in some experiments. The relevance of these findings is discussed in relation to the existing commercial fishery.     

A redescription of Tontonia cornuta (Leegaard, 1915) comb. nov., a planktonic oligotrichous ciliate (Ciliophora: Oligotrichia) from the northern Pacific Ocean

A rare marine planktonic ciliate, Tontonia cornuta comb. nov. [formally Strombidium cornutum (Leegaard, 1915) Kahl, 1932], from the northern Pacific Ocean was morphologically and biometrically investigated based on observations of permanent preparations after protargol impregnation. This species is similar to Tontonia simplicidens but can be distinguished by its conspicuously large size, higher number of macronuclei, and large number of buccal adoral membranelles.     

Alexandrium diversaporum sp. nov., a new non-saxitoxin producing species: Phylogeny,morphology and sxtA genes

Species of the PST producing planktonic marine dinoflagellate genus Alexandrium have been intensively scrutinised, and it is therefore surprising that new taxa can still be found. Here we report a new species, Alexandrium diversaporum nov. sp., isolated from spherical cysts found at two sites in Tasmania, Australia. This species differs in its morphology from all previously reported Alexandrium species, possessing a unique combination of morphological features: the presence of 2 size classes of thecal pores on the cell surface, a medium cell size, the size and shape of the 6″, 1′, 2⁗ and Sp plates, the lack of a ventral pore, a lack of anterior and posterior connecting pores, and a lack of chain formation. We determined the relationship of the two strains to other species of Alexandrium based on an alignment of concatenated SSU-ITS1, 5.8S, ITS2 and partial LSU ribosomal RNA sequences, and found A. diversaporum to be a sister group to Alexandrium leei with high support. A. leei shares several morphological features, including the relative size and shapes of the 6″, 1′, 2⁗ and Sp plates and the fact that some strains of A. leei have two size classes of thecal pores. We examined A. diversaporum strains for saxitoxin production and found them to be non-toxic. The species lacked sequences for the domain A4 of sxtA, as has been previously found for non-saxitoxin producing species of Alexandrium.     

Landscapes with high amounts of mass-flowering fruit crops reduce the reproduction of two solitary bees

The intensification of agriculture and the loss of natural habitat are key drivers of the observed declines in wild bee populations in recent decades. Mass-flowering crops are suggested to provide abundant resources for bees during flowering. However, mass-flowering crops could influence the offspring reproduction of bees after mass-flowering due to a lack of floral resources in the fields after mass-flowering. In our study we quantified the reproductive output of two solitary bee species (Osmia cornuta and O. bicornis) during and after mass-flowering of fruit crops (MFFC, i.e. apple, pear and sweet cherry) by means of trap nests. Ten trap nests were placed in landscapes with independent gradients of semi-natural habitat and MFFC in the surrounding landscapes. In addition, the composition of the collected pollen of both bee species was identified. Despite our relatively small sample size, we do detect a clear effect for both bee species, namely that the number of constructed brood cells after mass-flowering decreased significantly as the proportion of MFFC within 200 m of the surrounding landscape increased. During mass-flowering no effects were found, probably because O. cornuta collects significant amounts of Prunus spp., Malus spp. and Pyrus spp. pollen and because O. bicornis collects Malus spp. pollen to a certain extent. Yet, the negative effect after mass-flowering is clearly limiting the reproduction of O. bicornis, a solitary bee species with a later phenology compared to O. cornuta. The latter might also be a concern for other bee species that are active in late spring or summer. Landscape management needs to focus on providing additional floral resources to sustain viable populations of solitary bees and other pollinating insects after mass-flowering in landscapes with high amounts of MFFC.     

Mean individual dry weight and length-weight regressions of some zooplankton of Lake Kariba

Individual dry weights of various crustacean zooplankton in Lake Kariba were determined. All the zooplankters were collected from pelagic waters. Regression equations relating dry weight to body length were computed for those species or groups in which size classes could be separated: Bosmina longirostris, Ceriodaphnia cornuta, and copepod copepodites and adults. The mean individual dry weights obtained for most of these zooplankters are similar to those reported in other studies.