Spatial and temporal distributions of juvenile horseshoe crabs (Arthropoda: Chelicerata) approaching extirpation along the northwestern shoreline of the New Territories of Hong Kong SAR,China

Juvenile horseshoe crabs, Tachypleus tridentatus and Carcinoscorpius rotundicauda, with prosomal widths of between 11.5 and 102.5 mm, occur along Hong Kong's northwestern shoreline abutting Deep Bay where dissolved oxygen (DO) has been identified as the principal factor determining distribution and survival. Unreported upon hitherto, however, Tachypleus tridentatus occurred consistently in association with sea-grass (Halophila beccarii) beds, with DO concentrations of 8–14 mg·L^?1, a median grain size of >180 μm, interstitial water contents of <36% (sediment wet weight) and organic contents of <3.2% (sediment dry weight). Most individuals occurred between 60 and 120 m down from the shoreline although there appeared to be a trend of down-shore migration in summer, possibly to alleviate thermal stress. No individuals were found at sediment temperatures ≤20°C, i.e. in sub-tropical Hong Kong's winter, when they adopt buried repose. Abundances of emerged T. tridentatus, peaking in November, were synchronized with interstitial salinities of 22–26‰ and averaged 1–2 individuals per 100 m². Mean population density of T. tridentatus was 4–9 individuals per 100 m². Only four individuals of C. rotundicauda were found in August and September, suggesting a recent dramatic decline in numbers. The natal crèche environment for juvenile horseshoe crabs in Hong Kong is deteriorating and threatening their survival.     

How survival and food intake of tri-spine horseshoe crabs,Tachypleus tridentatus respond to thermal variation: implications for understanding its distribution limit

ABSTRACT

Temperature appears to have pervasive effects on larval development, feeding and movement patterns of tri-spine horseshoe crabs, Tachypleus tridentatus. To investigate how temperature determines their geographic distribution range, we examined the survival rate (SR), survival time (ST) and food intake of subadult T. tridentatus under different temperature levels in a seven-day experiment, followed by a three-day recovery to room temperature at 25°C. Significantly lower SR and ST of subadults were found at 40°C after the end of the seven-day experiment. Meanwhile, SR and ST of individuals at 0°C were negatively affected when the temperature was brought back to 25°C. Mean food intake of the subadults was statistically lower at 0–20°C and 35–40°C compared to that at 25°C and 30°C. After the subsequent three-day recovery to 25°C, only the individuals previously exposed to 15°C and 20°C had recovered to their normal level of feeding activities, but not the other treatment groups. These findings may provide preliminary data on how thermal tolerance determines the present distribution of T. tridentatus, which are found in large numbers around Beibu Gulf (annual temperature variation: 22–30°C), while considerably lower population densities are observed in the colder southern Sea of Japan and warmer seas of Southeast Asia.     

Life cycle and population structure of the terrestrial isopod Hemilepistus klugii (Brandt, 1833) (Isopoda: Oniscidea) in Iran

The life cycle and population structure of Hemilepistus klugii were studied in a population in Varamin, Iran. The population was sampled monthly (or fortnightly during the breeding season) from February 2008 to June 2009 and a total of 7015 individuals, comprising 1069 males, 1079 females and 4867 juveniles, were collected. As in other Hemilepistus species, five distinct phenophases, namely pair formation, gestation, hatching, growth and stationary, were recorded during the life cycle of H. klugii. The overall sex ratio was 1 : 1 but varied over time. Ovigerous females were observed only in April, indicating a seasonal and very short breeding period. With a short lifespan after breeding, females demonstrated true semelparity. The mean cephalothorax width for ovigerous/post-ovigerous females was higher in 2008 than in 2009. These females attained the largest size in the population throughout the year. The number of eggs per female ranged from 28 to 147 (mean ± SE, 78 ± 1.8). There was a positive correlation between female size and fecundity. Recruitment occurred in late April and resulted in the highest population density in this month, whereas the lowest densities were observed during November to January. Despite a high percentage of ovigerous females carrying undeveloped eggs (72.3%), intramarsupial mortality was low (3.5%).     

Carapace ontogeny of the bromeliad dwelling ostracod Elpidium bromeliarum Müller, 1880 (Crustacea: Ostracoda)

Ostracods have discontinuous development, marked by ecdyses, and determinate growth. Ontogenetic studies have suggested a pattern of nine postembryonic stages (eight juvenile phases and the adult) for the main freshwater ostracod superfamilies Cypridoidea, Cytheroidea and Darwinuloidea. However, cases of a supposed extra moult in the adult stage have been proposed for some ostracod species, which would amount to 10 postembryonic growth stages. One such example is Elpidium bromeliarum Müller, 1880, a cytheroidean that inhabits tank-bromeliads. The present study is aimed at investigating the ontogenetic development of E. bromeliarum using width and length measurements, as well as carapace and appendage morphology in order to test the existing hypothesis of an additional growth stage in adults of this species. Our results revealed nine postembryonic growth stages (eight juvenile and the adult), with sexual dimorphism beginning its expression in the last juvenile stage. Thus, the ontogenetic development of E. bromeliarum agrees with the overall pattern observed for podocopid ostracods and the hypothesis of 10 postembryonic growth stages was not corroborated. We argue that the inability to differentiate juveniles from adults, or different species from one another, may have misled to the assumption of an additional moult in E. bromeliarum.     

Biology of a tropical intertidal population of Cerithium atratum (Born, 1778) (Mollusca,Gastropoda)

The spatial distribution, population structure and secondary production of the gastropod Cerithium atratum were studied in the intertidal region of Engenho d'Água Beach in São Sebastião Channel on the northern coast of the state of São Paulo, Brazil, from May 1998 to April 1999. Twenty monthly samples were taken using a 0.25×0.25?m quadrat in each of two areas. Individuals showed a clustered dispersion pattern in both areas throughout the sampling period. Abundance and mean shell size were related to period of the year and to intertidal level, although no consistent pattern could be described when comparing the two sampling areas due to significant interaction between time and locality. Migratory events were evidenced by marked year-round variations in the frequency of individuals in the largest size classes. Thus, the size structure of this population could not be explained by recruitment and growth alone. Almost 90% of the intertidal population was represented by small individuals that recruited at the beginning of the sampling period (May to August 1998). The intertidal population was composed of annual cohorts formed by individuals recruited in different years. Each annual cohort could be divided into sub-cohorts with closely spaced recruitment periods in the same breeding season. The growth and annual P/B ¯ ratio (where P is the annual somatic production and B ¯ is the mean annual biomass) was estimated for the first year of life of recruits from 1998 occupying the intertidal region. Individuals may have an increment of up to 15?mm in shell length during the first year of life, but the growth was more pronounced in summer months. The high value of P/B ¯ (3.902) was a consequence of the dominance of small-sized individuals, which have a higher weight-specific increment than large-sized ones.     

The diets of juvenile horseshoe crabs,Tachypleus tridentatus and Carcinoscorpius rotundicauda (Xiphosura), from nursery beaches proposed for conservation in Hong Kong

Once common as a fishery resource, adult horseshoe crabs of two species occur in Hong Kong. A third, Tachypleus gigas, can no longer be found. Once common too were breeding and nursery beaches for horseshoe crabs in Hong Kong but overfishing of adults, pollution and coastal reclamation have reduced these to but three identified sandy mudflats and only one where juvenile Tachypleus tridentatus and Carcinoscorpius rotundicauda are sympatric. Nothing is known, however, of the conservation requirements of these two species locally and this study of the diets of these juvenile horseshoe crabs on two such nursery beaches aimed at providing information on this most elementary aspect of their biology. Gut contents of juvenile horseshoe crabs, i.e. nine Tachypleus tridentatus (8.5–67?mm prosoma width) and two Carcinoscorpius rotundicauda (50 and 52?mm prosoma width), were analysed and compared with the composition of the ambient assemblage of meiobenthos on a nursery beach at Pak Nai, Hong Kong. The obtained data suggest that juvenile horseshoe crabs of both species are selective benthic feeders and subsist mainly on insect larvae, polychaetes, oligochaetes, small crabs and thin-shelled bivalves. A strong, positive, preference for insect larvae (Chironomous sp.: Diptera: Chironomidae) was recorded, but with no preference for meiofauna over macrofauna.     

Aspects of the biology of a population of the cosmopolitan brittlestar Amphipholis squamata (Echinodermata) from the Firth of Forth,Scotland

In the Firth of Forth, Amphipholis squamata has continuous brooding, and pulsed breeding and recruitment, with highest numbers of brooded embryos and newly released juveniles occurring in the summer months. The estimated size of maturity was 1·5 mm disc diameter although the majority of reproductive output came from individuals of between 2·00–2·99 mm disc diameter. Brittlestar size and season had significant effects on embryo production. Embryos of different developmental stages were present in the bursae of the same individual, and newly spawned embryos and those of a potentially releasable size were present each month. The majority of juveniles apparently emerge at a disc diameter of between 0·8–1·2 mm.

Brittlestars were found with full stomachs throughout the year but a peak of feeding was recorded during summer when growth and seawater temperatures were highest. Stomach analyses indicated that almost all animals (99·4%) ingest fine particulate matter but a relatively wide range of plant and animal food items was also identified. Approximately 3% of this Scottish population of A. squamata was infested by the ectoparasitic copepod Cancerilla tubulata.     

The morphological differentiation of two horseshoe crab species,Tachypleus tridentatus and Carcinoscorpius rotundicauda (Xiphosura), in Hong Kong with a regional Asian comparison

Two horseshoe crab species, namely Tachypleus tridentatus and Carcinoscorpius rotundicauda, have been identified from Hong Kong. This paper describes the morphological characteristics of the two species and the morphometrics employed to help distinguish them. Comparisons were also made between adults from local waters and those from other Asian geographic regions. Intra-specific variations were also investigated by comparing morphological data obtained for local T. tridentatus juveniles and individuals of comparable sizes from Fujian Province, China. Morphological differences between the species and sexes and significantly different morphometric ratios were identified for the two horseshoe crab species. Intra-specific variations in morphometric characteristics were also demonstrated between juveniles from local beaches and those from Fujian.     

Mating behaviour,nympho-imaginal development and description of a new Mesabolivar species (Araneae: Pholcidae) from the Brazilian dry forest

A new species of Mesabolivar is described from Brazilian forests: Mesabolivar delclaroi. The mating behaviour, postembryonic development, number of egg sacs and eggs, birth rate, number of instars, developmental time until adulthood, sex ratio and size of cephalothorax (per instar) were recorded. The sexual behaviour was described and categorized into four steps: courtship, pre-copulation, copulation and post-copulation. After hatching, individuals presented five instars until maturity. The mean number of eggs (42 ± 16.6) and live births (31.5 ± 3.4) of the first egg sac were significantly greater than that of a second one (23.8 ± 3.8, and 19.25 ± 3.9, respectively). The developmental time from birth to adulthood (130.8 ± 9.6 days) did not differ significantly between egg sacs produced (128.61 ± 11.1). The size of the cephalothorax did not differ among adults or between sexes. The sex ratio revealed a shift in favour of females (4: 3).     

Breeding biology and population structure of the intertidal crab Petrolisthes laevigatus (Anomura: Porcellanidae) in central-southern Chile

The present study describes the seasonal reproductive cycle and population structure of the porcellanid crab Petrolisthes laevigatus, a typical inhabitant of the rocky intertidal. A total of 960 individuals (including 372 females) was collected in monthly samples from October 1993 to December 1994 in central-southern Chile. Ovigerous females were present between August and December, and were absent between January and July. The breeding season started during late winter (August), and females with embryos close to hatching occurred between October and end of spring (December). Considering the gonadosomatic index and the duration of the incubation period, it is concluded that P. laevigatus has a markedly seasonal breeding period with probably two successive spawnings per female per season. Our results compared with previous studies in temperate intertidal porcellanids suggest a lower structural variability of the population during the sampling period, but a pronounced seasonal variation in both sex ratio and abundance. The influence of environmental habitat conditions (i.e. temperature) is suggested to be responsible for the lower variation in the crab population size structure in ovigerous females, and the almost constant size distribution observed in males.     

Comparative population biology of Uca rapax (Smith, 1870) (Brachyura,Ocypodidae) from two subtropical mangrove habitats on the Brazilian coast

The structure of two populations of the fiddler crab Uca rapax in two subtropical mangrove habitats near Ubatuba, State of São Paulo, Brazil were compared. The size–frequency distribution, sex ratio, and recruitment were evaluated. Sampling was performed monthly from April 2001 to March 2002 in the Itamambuca and Ubatumirim habitats. Crabs were caught manually for 15?min by two collectors during low tide. The carapace width of each crab was measured with a digital caliper, and the sex and ovigerous state were recorded. The median size of the carapace width of males was greater than that of females at both sites (P<0.05). The median size of the crabs from Itamambuca was larger than at Ubatumirim (P<0.05). Only 28 ovigerous females were obtained from both mangroves, which suggested that females might remain in their burrows during the incubation period. The highest recruitment pulse occurred in winter for both populations, probably as a consequence of high reproductive activity during summer. The sex ratio in the size classes showed an anomalous pattern, with a higher frequency of females in the intermediate size classes. This may be related to a greater energy requirement for reproduction in females, thus delaying growth. The variable environmental conditions to which Uca rapax populations are subject appear to act directly or indirectly on the population, causing variations in growth and reproductive processes in the different populations investigated here.     

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.     

Population dynamics and phenology of two congeneric and sympatric lynx spiders Peucetia rubrolineata Keyserling, 1877 and Peucetia flava Keyserling, 1877 (Oxyopidae)

Numerous phenological studies on spiders belonging to the families Oxyopidae, Thomisidae, Lycosidae, Selenopidae (e.g. Selenops cocheleti) and Salticidae (e.g. Psecas chapoda and Psecas viridipurpureus) have been conducted in the neotropical region. However, studies that simultaneously compare population dynamics and age structure in populations of sympatric species are limited, especially in Oxyopidae. The population dynamics and phenology of two congeneric lynx spider (Peucetia rubrolineata and Peucetia flava) were examined in southeastern Brazil. Several characteristics of the age structure were compared between the two spider species. The variation in the total abundance of individuals and age structure and their relationship with climatic variables were similar between the two spiders. Adults of these spiders were present mainly in spring, indicating an annual reproductive cycle and a ‘stenochronous spring’ phenological pattern. The recruitment of spiders occurred in summer for P. rubrolineata and P. flava, followed by successive phenological peaks at all stages of development. Rainfall and temperature had a positive correlation to population flux in the two spider species studied. Despite the significant climatic effects observed in the Serra do Japi, the phenological pattern of the population in this study was not always repeated in other species of spiders (e.g. Selenops cocheleti).     

Microdistribution of juveniles and adults of the mud crab Panopeus americanus (Brachyura,Panopeidae) in a remnant mangrove area in the southwest Atlantic

The distribution and density of a population of the narrowback mud crab Panopeus americanus were evaluated, with emphasis on the different developmental stages (juveniles and adults) and sexes. Specimens were collected in a remnant human‐impacted mangrove at Araçá on the northern coast of São Paulo, Brazil, at two‐month intervals from January through July 2006, using 1 m×1 m sample plots randomly distributed in the intertidal zone. The population was abundant, with a mean density of 14.3±9.3 individuals/m². Larger individuals were more abundant in the upper and middle intertidal zones, whereas smaller crabs were predominant in the lower intertidal and thus remained submerged longer. Males and juveniles were homogeneously distributed. Females preferred regions farther from the waterline. A clustered distribution pattern suggested gregarious behaviour and settling of juveniles in the vicinity of the adults' habitats, in an adaptive process as a function of tidal rhythms.     

Demography of the Yarrow's spiny lizard, Sceloporus jarrovii , from the central Chihuahuan Desert

The demography of a population of Yarrow's spiny lizard, Sceloporus jarrovii , was examined from 2004 to 2006 in the canyon Las Piedras Encimadas, located in Gomez Palacio, Durango, México. Lizards were studied using a mark-recapture technique. Reproduction in females occurred between November and May, coinciding with dry conditions. Reproductive activity was highest (percent of females with vitellogenic follicles or embryos) in the middle of the dry season (November and December). Thirteen percent of females reached sexual maturity at an average age of 8.5 months. The population structure was similar in spring and fall, but not in summer. A notable feature of summer, coinciding with the wet season, was the greater number of hatchlings and juveniles. The overall sex ratio did not differ from 1:1. The density of adults varied from 12 to 62 animals ? 0.5 ha –1 . Temperate and arid-adapted populations of S. jarrovii exhibited broad similarity in timing of the reproductive season, whereas factors such as density, growth, age at sexual maturity, and survivorship differed.     

Observations on mermithid parasitism (Nematoda: Mermithidae) in two species of Lasius ants (Hymenoptera: Formicidae)

The morphological effects of nematode parasitism by Pheromermis myrmecophila were investigated in two species of ants, Lasius flavus and Lasius niger, collected on Clare Island, Ireland. Infected males and queens were found for L. flavus but only infected queens in L. niger. The numbers of nematodes present in each infected specimen were counted and their lengths were measured. Head width, femur length and wing measurements were made for infected and non-infected individuals of both species. In queens, the length of the nematodes decreased as the number per ant increased. When single nematodes were present their mean lengths were significantly greater in L. flavus than in L. niger. In addition to the reduction in wing size for all parasitized specimens, head widths and femur lengths of the queens were significantly different in both species. A log–log graph of radial cell length versus width suggested a linear reduction of wing size.     

Estimating population features of the anomuran crab Petrolisthes armatus (Porcellanidae) in a remaining and impacted mangrove area of the western Atlantic

This study evaluated biological aspects of Petrolisthes armatus inhabiting a remaining but disturbed mangrove area of Brazil. Samples were taken from March 2005 to July 2006, during low tide in the rocky-shore region. The size frequency distribution for all the individuals collected was bimodal. The sex ratio obtained was not different from 1:1, and the population was classified as standard. A higher reproductive ratio and earlier ovigerous conditions were found in relation to another southern population; these can be hypothesized as adaptations to life in a stressed environment. Even though the population living in Araçá has been subject to an environment frequently disturbed by human-produced pollutants, our results show no sign of negative effects on reproductive stages, recruits or members of the population in general. The population profiles of P. armatus show some peculiarities when compared to other populations inhabiting non disturbed environments.     

Population dynamics of the bark-dwelling spider Eustala perfida Mello-Leitão, 1947 (Araneidae)

Population dynamics of spiders may vary according to biotic and abiotic factors. We studied the phenology of the bark-dwelling spider Eustala perfida over a two-year period and investigated how temperature, precipitation, and prey availability were associated with the population of this species. The population of E. perfida presented a well-defined fluctuation, with one reproductive cycle per year. The recruitment of spiders occurred in summer, followed by successive phenological peaks at all stages of development. The species is protogynic, and the population of adult males peaks in the cold/dry season, indicating a winter mature phenological pattern. The diet of E. perfida is composed mainly of medium-sized homopterans, complemented by small dipterans and hymenopterans. We found that the higher abundance of spiders occurred two months after the highest values of precipitation and temperature, and three months after the higher abundance of potential prey items. These results indicate that spiders require time to respond to changes in environmental conditions. The species’ natural history and the structure of the trunks in which it lives may also affect the number of individuals in a particular area and over time.     

Reproductive strategies of two sympatric species of Hyalella Smith, 1874 (Amphipoda,Dogielinotidae) in laboratory conditions

This study was performed in order to test theoretical predictions that sympatric species might show different reproductive strategies to facilitate their coexistence. The duration of precopulatory mate‐guarding behaviour, body size of ovigerous females, embryonic and postembryonic development time, fecundity, and viability of the sympatric amphipod species Hyalella pleoacuta and H. castroi were investigated under laboratory conditions (19±1°C and 12 h light/dark photoperiod). Specimens were collected with nets in autumn and spring 2004, and summer and winter 2005 in the Vale das Trutas, São José dos Ausentes County, southern Brazil, and transported to the laboratory. Both species of Hyalella showed similar reproductive strategies, especially in respect to the duration of the precopula, duration of the embryonic period, and fecundity. However, these species differed in body size of ovigerous females, duration of the postembryonic period, viability, and body size of juveniles that remained in the marsupium and those released by females. These differences in the reproductive traits of H. pleoacuta and H. castroi may facilitate their coexistence in nature.     

Population and reproductive biology of the mud crab,Macrophthalmus hirtipes (Jacquinot, 1853) (Ocypodidae), from marine and estuarine habitats

Summary

Monthly samples of the burrowing mud crab, Macrophthalmus hirtipes (Jacquinot 1853) were collected from February 1978 to April 1979 from a marine inlet, Governors Bay (43°38′S, 172°39′E) and from the Avon-Heathcote Estuary (43°33′S, 172°44′E), New Zealand. The estuarine population was generally unimodal and had an annual cycle of population change sustained by a concentrated summer recruitment. In contrast, the bay population was unimodal and bimodal at different times, and recruitment occurred thoughout the year with no clear peak. Both populations had equal sex ratios and were dominated by immature individuals (crabs?10·0 mm carapace width). Estuarine crabs were generally smaller than marine crabs.

At both sites, ovigerous females were found from late May to early March; the smallest egg-bearing female measured CW 10·0 mm at the estuary and CW 10·5 mm at the bay. Cycles of ovarian and embryological development were similar at the two localities. Three peaks of egg oviposition occurred within one breeding season and mature females were capable of producing successive broods. Fecundity was related exponentially to female carapace width and did not alter during the breeding season. Estuarine females carried more eggs than marine females of similar size. Egg volume was related to time of oviposition and eggs laid at the start of the breeding season were significantly larger than those deposited during the second half; however, estuarine eggs showed less reduction in size at this time than did marine eggs.

Estuarine sediments had higher levels of chlorophyll ‘a’, ATP, and microorganism activity and productivity than marine sediments. As M. hirtipes is a detritus feeder, the differences in food resources are believed to account for the elevated fecundity of estuarine crabs. At the bay, salinities were high with little tidal and seasonal variation, whereas in the estuary sea water was diluted significantly both daily and seasonally. The differences in salinity characteristics between the sites may be affecting larval distribution; recruitment in the estuary was concentrated to the summer and in the bay it was almost continuous.