Larval development of the pedunculate barnacles Octolasmis angulata Aurivillius 1894 and Octolasmis cor Aurivillius 1892 (Cirripedia: Thoracica: Poecilasmatidae) from the gills of the mud crab,Scylla tranquebarica Fabricius, 1798

Detailed studies of larval development of Octolasmis angulata and Octolasmis cor are pivotal in understanding the larval morphological evolution as well as enhancing the functional ecology. Six planktotrophic naupliar stages and one non-feeding cyprid stage are documented in details for the first time for the two species of Octolasmis. Morphologically, the larvae of O. angulata and O. cor are similar in body size, setation patterns on the naupliar appendages, labrum, dorsal setae-pores, frontal horns, cyprid carapace, fronto-lateral gland pores, and lattice organs. Numbers of peculiarities were observed on the gnathobases of the antennae and mandible throughout the naupliar life-cycle. The setation pattern on the naupliar appendages are classified based on the segmentation on the naupliar appendages. The nauplius VI of both species undergoes a conspicuous change before metamorphosis into cyprid stage. The cyprid structures begin to form and modify beneath the naupliar body towards the end of stage VI. This study emphasises the importance of the pedunculate barnacle larval developmental studies not only to comprehend the larval morphological evolution but also to fill in the gaps in understanding the modification of the naupliar structures to adapt into the cyprid life-style.     

A new species of Tigriopus (Copepoda, Harpacticoida, Harpacticidae) from Thailand with the description of its naupliar development

Both genders of Tigriopus thailandensis sp. nov. are described from a laboratory stock raised from individuals collected from the seaweed Enteromorpha clathrata in Thailand (Bangsaen Beach, Chonburi Province). Tigriopus thailandensis sp. nov. shares with its closest relative T. japonicus Mori, 1932 two setae on the third exopodal segment of leg 4 while other congeners bear 3 inner setae. However, allobasis and exopod of antenna in both genders are much more slender and elongate than in T. japonicus. All six naupliar stages of T. thailandensis are described from the offspring of isolated females. In comparison with nauplii of T. japonicus, T. thailandensis nauplii are characterized by the following: a smaller body size throughout the naupliar phase; first antennular segment without seta, second antennular segment with only one small seta plus two longer setae; third antennular segment with additional spinules from naupliar stage II onwards; antenna bears three small spinules on the terminal exopodal segment; one additional seta on the anterior surface of the antennary basis, tubular endopod of antenna with one tiny seta midlength at naupliar stage III that increases in size; mandibular basis with several spinules on anterior surface; mandibular coxa with one spinulose seta that is smooth in T. japonicus.     

Growth and development of Calanus chilensis nauplii reared under laboratory conditions: testing the effects of temperature and food resources

We assessed growth and development of naupliar stages of Calanus chilensis Brodsky 1959, under a combination of three temperatures and two food levels in laboratory conditions. Both food supply and temperature significantly affected naupliar growth and development. High food, measured as chlorophyll a (Chl a) concentration, was 40 μg l⁻¹, on average, and yielded temperature-dependent growth rates in the range of 0.13-0.17 day⁻¹. Low food was about 1.2 μg chlorophyll a l⁻¹ and retarded or arrested development and drastically reduced the growth rate to the range of 0.05-0.09 day⁻¹. To test whether these experimental results were consistent with field data, we used published information on temperature and chlorophyll a variability in northern Chile and developed a combined temperature/food-dependent model to diagnose naupliar growth in the field through a 2-year seasonal cycle including the 1997-1998 El Niño conditions. We concluded that in the upwelling region off northern Chile, C. chilensis might seldom encounter conditions of food shortage, as those applied in the laboratory. Thus, naupliar growth of this species may be primarily controlled by environmental temperature and this might also be the case for the dynamics of the entire population inhabiting the coastal upwelling zone.     

The larval development of an Australian limnadiid clam shrimp (Crustacea, Branchiopoda, Spinicaudata), and a comparison with other Limnadiidae

The adult morphology of the Australian Limnadopsis shows some remarkable differences to that of other Limnadiidae. These differences are not reflected in its larval development. In Limnadopsis parvispinus, larval development comprises six stages. In stages I and II only the three naupliar appendages are present: the antennule as a small bud, the biramous antenna as the main swimming organ, and the mandible. The antennal protopod bears two endites, the proximal naupliar process and a more distal endite. In stage III a bifid naupliar process (in earlier stages not bifid) and the first signs of the carapace and trunk limb anlagen (undifferentiated rudiments) appear. In stage IV the carapace anlagen become more pronounced. The number of trunk limb anlagens increases to five, and differentiation has commenced. In stage V the first five pairs of trunk limbs are differentiated to varying degrees. The anterior-most four pairs of trunk limbs are subdivided into five endites, a small endopod, an exopod and an epipod. The bivalved carapace covers the anterior-most limbs. In larval stage VI the carapace is larger and the trunk limbs are further differentiated. A general pattern in the sequence of larval stages is the increasing number of sensilla on the antennules. From the last larval to the first postlarval stage, a significant change in morphology takes place. The trunk limbs are now used for swimming. Typical larval organs are much smaller than in the last larval stage. A comparison with other representatives of the Limnadiidae shows a high degree of correspondence, with most differences explained by the heterochronous appearance of characters during development. Five to seven stages are described for all studied Limnadiidae, including one particular stage in which four fully developed setae, a bifid naupliar process and the first signs of carapace anlagen are present. These characters are found in stage III in L. parvispinus, Limnadia stanleyana, Eulimnadia texana, and Imnadia yeyetta but in stage IV in E. braueriana and L. lenticularis. Based on a comparison of the larval stages of six limnadiid and one cyzicid species, we conclude that at least six naupliar stages belong to the limnadiid ground pattern.     

Cannibalistic behaviour of rock-pool copepods: An experimental approach for space, food and kinship

Studies on cannibalism in harpacticoid copepods are restricted to predation on naupliar larvae in rock-pool harpacticoids of the genus Tigriopus. An earlier experimental study on the Mediterranean copepod Tigriopus fulvus indicated that females recognized their own larvae and preferentially preyed on nauplii other than their own. In a series of laboratory experiments, we tested if there were differences in naupliar predation as a function of crowding, food level and sex in Tigriopus brevicornis and T. fulvus. Results show that cannibalism was restricted to the first larval stages (N1 and N2). Both food availability and adult density significantly affected the predation rate. Contrary to earlier suggestions, adult males also preyed on the nauplii. We found no evidence that adults spare their own offspring, for neither T. fulvus nor T. brevicornis.This is in accordance with what one would expect for species having the life history characteristics of Tigriopus, i.e.: multiple broods and large number of offspring. Earlier results indicating parental care in Tigriopus must be taken with caution.     

Postembryonic development of Drescheriella glacialis Dahms & Dieckmann (Copepoda,Harpacticoida) reared in the laboratory

Summary All 6 naupliar and 5 copepodite stages of Drescheriella glacialis, a tisbid from Antarctic fast ice, are described. A key for the identification of the naupliar stages is given. Sexual dimorphism makes its first appearance in the copepodid IV. The setae configuration of the exopod of the second antenna in copepodid I is considered to be an ancestral state, and subsequent ontogenetic reductions are proposed to be apomorphic processes.     

Estimate of the effects of ultraviolet radiation on the mortality of Artemia franciscana in naupliar and adult stages

The impact of different doses of artificial ultraviolet (UV) radiation on the growth stages of a marine zooplankton was investigated using laboratory microcosms. Mortality percentages of naupliar and adult samples of Artemia franciscana were recorded in relation to different UV doses (single exposure: 75, 150, 300, 600, 1,200, 2,400, 3,900, 7,800 J m^–2) at specific observation times after exposure (24, 48, 72, 96 and 120 h). The relationship between mortality percentage and UV dose showed significant differences in relation to the zooplankton growth stage. The elevated susceptibility of the naupliar samples to UV radiation is described through a mortality model based on a logistic equation. The data analysis shows that the slope of mortality versus dose remains the same for the two growth stages while the lethal dose in the naupliar stage was 3.3 smaller than that determined for the adult stage. The slope of the UV mortality rate versus post-incubation time was found to be significantly different (P<0.05) at low UV doses for the two life stages examined, i.e. naupliar and adult. The lower value of LD₅₀ in naupliar stages compared to that for adults confirms that in the early growth stage this marine zooplankton is more susceptible to UV radiation.     

Ontogenetic niches of a planktonic copepod in Lake Biwa studied on a fine temporal scale

Eodiaptomus japonicus was collected in the north basin of Lake Biwa, Japan, on six dates from 11 to 25 June 1987. Temporal changes in its vertical distribution and reproduction indices were small, but those in its abundance were large. Mortality was high in the naupliar stage I and from copepodid stage III to adult stage. The latter fact and low proportion of females in adulthood suggest heavy predation by fishes which prefer larger prey.Eodiaptomus japonicus showed ontogenetic vertical migration within the epilimnion. The direction of the migration was upward in the early naupliar stages and downward in the late naupliar and early copepodid stages. Both migrations resulted in raising mortality in the corresponding or succeeding stages, probably expressing the behavioral constraints ofE. japonicus.     

Redescription of Mesocyclops leuckarti (Copepoda,Cyclopoida), Including a Study of its Naupliar Development

The female and male of Mesocyclops leuckarti (CLAUS , 1857) from a European population are redescribed in detail. New diagnostic characters are described which have passed unnoticed so far. All six naupliar stages of this species are studied as well. Characters of the adult and naupliar external morphology are discussed with respect to previous studies.     

The complete postembryonic development of Tropodiaptomus informis Kiefer, 1936 (Copepoda:Calanoida) reared in the laboratory

The complete postembryonic development of Tropodiaptomus informiscomprises six naupliar and six copepodid stages of which thelast is the adult. Of all the characteristics, it is the shapeand number of setae on the terminal segments of the antennulesin the late naupliar stages, and the shape and setae of exo-and endopods of the fifth legs from copepodids III to V whichare most useful in constructing relationships among eight diaptomidsreared.     

Development and mortality of the first naupliar stages of Eurytemora affinis (Copepoda, Calanoida) under different conditions of salinity and temperature

As a prevalent species complex in temperate estuaries and salt marshes of the Northern Hemisphere, populations of Eurytemora affinis that inhabit these environments must be adapted to salinity fluctuations. Some populations have invaded freshwater environments. In this work, we focus on the combined effects of temperature and salinity fluctuations on mortality rates and development time of the first naupliar stages under starvation. Two temperatures (10 and 15 °C) and eight salinities, ranging from 0 to 35 psu are investigated. We show (i) that among all experimental conditions the optimal temperature and salinity for naupliar survival and development are 15 psu and 15 °C, and (ii) that only the most extreme salinities (i.e. 0 and 35 psu) have a negative effect on naupliar survival. Nauplii develop faster and reach a higher developmental stage at 15 than at 10 °C, independent of salinity. The relevance of this metabolic adaptive pattern is discussed in the general framework of in situ behavior, tidal forcing and biogeographic variability, as well as the potential sources of the observed individual variability.     

Larval development of Octomeris sulcata Nilsson-Cantell (Cirripedia: Thoracica: Chthamalidae) from Japan and Korea

Embryos obtained from gravid adults of the chthamalid barnacle Octomeris sulcata Nilsson-Cantell from Japan and Korea were cultured through six naupliar stages to the cyprid and juvenile barnacle stage in laboratory conditions, fed either the diatom Skeletonema costatum (Grev.) Cleve or the dinoflagellate Prorocentrum minimum (Pavillard) Schiller. The nauplii were planktotrophic and, depending on diet, reached the cyprid stage 9 or 17 days after hatching in individual cultures at 22 °C with 24 h illumination. The survival rate was higher and the duration of the naupliar stages was shorter when fed P. minimum rather than S. costatum. This is probably due to the presence of feathered setae on the antennae. Feathered or plumose setae in nauplii of different cirripede taxa are apparently linked to the type of phytoplankton in the seas when these taxa first evolved.The larval stages of O. sulcata are described, and morphological differences between larvae reared from Japanese andKorean adults are compared. The polygonal cephalic shield and unilobed labrum, a pair of posterior shield spines after naupliar stage IV, feathered setae and a hispid seta on the coxa of the antenna, a cuspidate seta on the mandible, and the gnathobase of the antenna are important in distinguishing the nauplii of this species from other species, including Chthamalidae.     

Copepod post-embryonic durations: pattern,conformity, and predictability. The realities of isochronal and equiproportional development,and trends in the opepodid-naupliar duration ratio

A collation of post-embryonic durations for freshwater and marine calanoid, cyclopoid and harpacticoid copepods is provided, and examined for patterns and conformities which may be of predictive value. Most of the analysis concerns calanoids. Only the genus Acartia exhibits evidence of equal stage duration (isochronality). Accordingly, isochronal development must be rejected as a general pattern in copepods — with various implications to the study of their production. Conversely, relative stage durations are surprisingly comparable in a wide range of copepods across a broad range of temperatures and food levels. A loose consistency is evident among copepod species generally, (interspecific equiproportionality), but within given genera, a striking regularity, here termed intergeneric equiproportionality (IGE) is evident. Patterns of IGE are consistent with selectively adaptive life-history traits, and IGE thus offers predictive prospects which are of both quantitative and qualitative (heuristic) value. Empirical support which exists for IGE among marine calanoids suggests that in contrast to isochronality and interspecific equiproportionality, IGE is indeed a real, and ultimately quantifiable feature of copepod development. Within calanoids, the ratio of total copepodid to total naupliar duration (Dc/Dn) appears independent of temperature, shows little relationship to adult body mass, but is inversely related to food supply, markedly so in freshwater forms. In this context, changes in the ratio are attributable largely to influences of food supply upon copepodid development times: naupliar durations appear relatively independent of food concentrations during development. Considerably higher Dc/Dn ratios in freshwater than in marine calanoids conceivably arise from selective influences of different nutritional conditions and size-selective predation pressures in freshwater and marine environments. In the generally high predation environment of freshwaters, acceleration of naupliar development potentially reduces the vulnerability of these smaller stages to size-selective tactile predation, while larger copepodid instars are able to reduce the opposing size-selective influences of visual planktivores by virtue of their escape responses to suction attack. Within the spectrum of copepod life histories and development schedules, ecologically consistent trends of this nature await formal recognition. The present recognition of IGE is a modest initiative in this quest. Both naupliar and copepodid durations are inverse monotonic functions of temperature. Several mathematical expressions which account for this temperature-duration response are provided. The response envelope is much tighter for naupliar than copepodid durations. Predictability of temperature-duration responses accordingly declines ontogenetically from egg to naupliar to copepodid stages.     

Naupliar and Metanaupliar Development of Thysanoessa raschii (Malacostraca,Euphausiacea) from Godth?bsfjord,Greenland, with a Reinstatement of the Ancestral Status of the Free-Living Nauplius in Malacostracan Evolution

The presence of a characteristic crustacean larval type, the nauplius, in many crustacean taxa has often been considered one of the few uniting characters of the Crustacea. Within Malacostraca, the largest crustacean group, nauplii are only present in two taxa, Euphauciacea (krill) and Decapoda Dendrobranchiata. The presence of nauplii in these two taxa has traditionally been considered a retained primitive characteristic, but free-living nauplii have also been suggested to have reappeared a couple of times from direct developing ancestors during malacostracan evolution. Based on a re-study of Thysanoessa raschii (Euphausiacea) using preserved material collected in Greenland, we readdress this important controversy in crustacean evolution, and, in the process, redescribe the naupliar and metanaupliar development of T. raschii. In contrast to most previous studies of euphausiid development, we recognize three (not two) naupliar (= ortho-naupliar) stages (N1-N3) followed by a metanauplius (MN). While there are many morphological changes between nauplius 1 and 2 (e.g., appearance of long caudal setae), the changes between nauplius 2 and 3 are few but distinct. They involve the size of some caudal spines (largest in N3) and the setation of the antennal endopod (an extra seta in N3). A wider comparison between free-living nauplii of both Malacostraca and non-Malacostraca revealed similarities between nauplii in many taxa both at the general level (e.g., the gradual development and number of appendages) and at the more detailed level (e.g., unclear segmentation of naupliar appendages, caudal setation, presence of frontal filaments). We recognize these similarities as homologies and therefore suggest that free-living nauplii were part of the ancestral malacostracan type of development. The derived morphology (e.g., lack of feeding structures, no fully formed gut, high content of yolk) of both euphausiid and dendrobranchiate nauplii is evidently related to their non-feeding (lecithotrophic) status.     

Cannibalistic feeding behavior of the brackish-water copepod Sinocalanus tenellus

Cannibalistic feeding behavior of the brackish-water copepodSinocalanus tenellus was examined in the laboratory using CI-II,CIII-IV and CVI female as predators and NI-II, NIII-IV, NV-VIand CI-II as prey. In each prey-predator combination, the ingestionrate increased with increasing prey density to an asymptoticvalue. Cannibalism took place even when phytoplankton was availableas an alternative food supply. Based on a daily ration, theoptimal prey stages for CVI females, CIII-IV and CI-II are NI-VI,NI-IV and NI-II respectively. Under average, natural prey density(10 nauplii l^–1), S tenellus can achieve only a smallfraction (max 9%) of the daily minimum food requirement by cannibalisticfeeding. However, the impact of cannibalism on naupliar survivorshipcan be significant. When adult females occur at a density of10 l^–1, the mortality due to cannibalism attains 99.2%during the naupliar stages.     

The Unique Dorsal Brood Pouch of Thermosbaenacea (Crustacea,Malacostraca) and Description of an Advanced Developmental Stage of Tulumella unidens from the Yucatan Peninsula (Mexico), with a Discussion of Mouth Part Homologies to Other Malacostraca

The Thermosbaenacea, a small taxon of crustaceans inhabiting subterranean waters, are unique among malacostracans as they brood their offspring dorsally under the carapace. This habit is of evolutionary interest but the last detailed report on thermosbaenacean development is more than 40 years old. Here we provide new observations on an ovigerous female of Tulumella unidens with advanced developmental stages in its brood chamber collected from an anchialine cave at the Yucatan Peninsula, which is only the third report on developmental stages of Thermosbaenacea and the first for the genus Tulumella. Significant in a wider crustacean context, we report and discuss hitherto unexplored lobate structures inside the brood chamber of the female originating at the first (maxilliped) and second thoracic segments, which are most likely modified epipods, perhaps serving as gills. At the posterior margin of carapace of the female are rows of large spines preventing the developing stages from falling out. The external morphology of the advanced developmental stages is described in much detail, providing information on e.g., carapace formation and early limb morphology. Among the hitherto unknown structures in the advanced developmental stages provided by this study are the presence of an embryonic dorsal organ and rudimentary ‘naupliar processes’ of the second antennae. Since most hypotheses on crustacean (and malacostracan and peracaridan) relationship rest on external limb morphology, we use early limb bud morphology of Tulumella to better establish thermosbaenacean limb homologies to those of other crustaceans, which is a necessary basis for future morphology based phylogenetic considerations.     

Effect of culture density and antioxidants on naupliar production and gene expression of the cyclopoid copepod, Paracyclopina nana

Although attempts have been made to use mass cultures of marine copepods as live foods in marine aquaculture, some limitations such as low density culture still exist. The brackish water cyclopoid copepod, Paracyclopina nana has the potential for mass culturing as live food. In this study, we not only investigated the effect of culture density on the naupliar production and specific gene expressions of P. nana, but also the effect of several antioxidants under the conditions of a high density culture. The naupliar production of the copepod decreased with increasing culture density. The expression of glutathione reductase (GR), selenium-dependent glutathione peroxidase (SeGPx), glutathione S-transferase kappa (GST kappa), heat shock protein 40 (Hsp40), and Hsp70 genes of P. nana increased in the high density treatment but vitellogenin genes (Vg1 and Vg2) showed downregulation. In the condition with 20 inds./mL, vitamin C had a significant decrease but sodium selenite induced the naupliar production of P. nana greatly. The expressions of GR, SeGPx, Hsp70, and Vg genes increased with the vitamin C treatment. Sodium selenite caused a decrease of SeGPx and Hsp40 but GST kappa increased in the treatment with 20 inds./mL. These results suggest that sodium selenite is a positive antioxidant which can increase the culture efficiency of the copepod.     

The development of the nervous system in Laevicaudata (Crustacea, Branchiopoda): insights into the evolution and homologies of branchiopod limbs and ‘frontal organs’

We investigated the development of the external morphology and of the nervous system in Lynceus biformis and Lynceus brachyurus (Laevicaudata, Branchiopoda), by using immunohistochemical methods in combination with a confocal laser scanning analysis. In both Lynceus species, a free-swimming nauplius larva, equipped with three appendages, hatches from resting eggs. Despite their close phylogenetic relationship to each other, considerable differences are present in their external morphology. Hatching L. brachyurus larvae are equipped with a large and flattened labrum, where in contrast, the L. biformis larvae possess a smaller labrum with four conspicuous posteriorly directed spines at its margin. Despite these differences, the development of the nervous system is quite similar in both species. The hatching larvae are equipped with a naupliar nervous system, and only in the more advanced stages, the development of the ventral nerve cord starts. Furthermore, our investigation into the nervous system provided insights into architecture and evolution of protocerebral sensory organs, the dorsal setae field and the dorsal frontal organ, only present in Laevicaudata. The identification of frontal filaments with an associated frontal filament organ in Lynceus revealed—after a comprehensive comparison with other branchiopods—that these organs exist throughout Branchiopoda and are comparable to those in other crustaceans. Additionally, our results of the peripheral nervous system analysis showed that the innervation pattern of the naupliar appendages (antenna and mandible) and the trunk appendages could be serially homologized, despite much difference in gross morphology of these. Based on the innervation pattern of limbs, we suggest that the larval uniramous mandibular palp, found in the larvae of all ‘large’ branchiopods, is largely exopodal of nature (contrary to most earlier statements) and that the endopodite of the trunk limbs consists of only one distal endite-like segment (confirming some earlier statements) and not of three as proposed by others.     

The naupliar stages of Cyclopina yutimaete Lotufo (Cyclopinidae,Cyclopoida)

The six naupliar stages of Cyclopina yutimaete Lotufo are described and compared to other cyclopoid nauplii. They seem more similar to freshwater cyclopoid nauplii than to marine species. The hirsute aspect of the last segment of the antennule is of diagnostic value for these nauplii.