Oldest shrimp and associated phyllocarid from the Lower Devonian of northern Russia

A mass occurrence of schooling, weakly sclerotized eocaridid shrimps in the Early Devonian pelagic environment of northern Russia suggests that the body-plan of the Eumalacostraca originated as an adaptation to swimming in the open sea. They probably replaced their ecological analogues, the archaeostracan phyllocarids Caryocaris , as a consequence of the rebuilding of the global marine environment during the later Ordovician. This may explain the rarity of the eumalacostracans in the Devonian and the difficulty with documenting their links with the archaeostracans. The new eocaridid shows similarities to the Carboniferous Anthracophausia in the general appearance and weak sclerotization of the carapace and appendages, but lacks its derived characters. The morphology of the pleural lobes, which have minute spines, is the only identified distinguishing character of the proposed new genus and species Archangeliphausia spinosa gen. et sp. nov. Its carapace was probably firmly connected dorsally with the thoracic segments. The only associated fossils of the assemblage represent a bizarre archaeostracan Pechoracaris aculicauda gen. et sp. nov., with a weakly sclerotized carapace lacking any hinge or rostral plates but with reduced furca, an extremely long caudal spine and pleopods transformed into spines.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 83–90.     

Ordovician phyllocarids (Arthropoda; Crustacea) from Argentina

New phyllocarids are described from the Ordovician (early Arenig to lower Caradoc) of the Precordillera and Eastern Cordillera of Argentina, namelyCaryocaris delicatus n. sp.,Caryocaris sp. 1, ?Caryocaris sp. 2,Pumilocaris granulosus n. gen., n. sp., ?Pumilocaris sp. and an undetermined form. They all occur in graptolite-bearing shales. A critical review of the South American phyllocarids formerly described in the literature is made andCaryocaris delicatus n. sp. (Llanvirn) is compared with congeneric species known from the Ordovician (Tremadoc-Caradoc) of North America, Europe, China and Australia. The presence ofCaryocaris in South America is therefore attested, which confirms the view that the genus had a worldwide distribution in the lower Ordovician in relation with a possible pelagic lifestyle. A new definition ofCaryocaris is proposed and the former subdivision of the genus [C. (Caryocaris) andC. (Rhinopterocarys)] abandoned. A set of original characters (e.g. lack of rostral plate, short leaflike furcal rami, short telson, extremely thin carapace) strongly indicates thatCaryocaris lies outside the Ceratiocarididae and may form the basis of a new taxonomic unit.     

A new Diacyclops (Copepoda,Cyclopoida, Cyclopidae) from northwestern Argentina

No species of the genus Diacyclops has been so far cited from continental Argentina. Specimens collected from a roadside pool in the province of Catamarca on a plateau at 3700 m a.s.l. turned out to belong to a new species, Diacyclops andinus, which is described and illustrated in detail for both sexes. Main diagnostic features are: spine formula of swimming legs 3, 4, 4, 4; terminal spines of the distal endopod segment of leg 4 subequal; smooth furcal rami. The new species is a member of the D. bicuspidatus group, and is probably near the base of this line.     

Variation in the species of freshwater harpacticoid copepods in Japan. II. Attheyella nakaii (Brehm)

Morphological variation in Attheyella nakaii, a dominant species of the genus in the mountain waters of Japan, is presented. The greatest proportion of male type furcal rami in the females of local populations is seen at the northern border of the range, where this species shows a sympatric distribution with A. yesoensis. Diminution of the inner lateral setae of endopodites on legs 2–4 occur in Kyushu and the Ryukyu Islands. The body size and relative length of the leg 5 exopodite show a latitudinal cline. The population of the type locality of this species exhibits a truncated form of furcal rami.     

Juvenile morphology of the xanthid crab Leptodius exaratus (H. Milne-Edwards, 1834) (Decapoda: Brachyura), with notes on the appearance of sexual dimorphism

Summary

The morphometrical and meristic features of the carapace, cephalic appendages (antenna, antennule), mouthparts (maxillule, maxilla, first-third maxillipeds), sternum, pereiopods, abdomen, and pleopods of juveniles and the onset of morphological sexual dimorphism were described for the xanthid crab Leptodius exaratus (H. Milne Edwards, 1834), based on laboratory-reared and wild adult specimens collected from Tateyama Bay, Japan. First instar juveniles shared some of the features of adults (e.g. gross appearance of the carapace and cheliped propodus proportions), but differed from adults on almost all other morphological parameter examined. Morphological development was still not complete at the ninth instar; extrapolation from the rate of morphological changes between instars 1–9 suggests that L. exaratus requires about 13 ecdyses to transform into adults, including development of reproductive structures. Differences in the number and morphology of pleopods and abdomen width allowed early distinction of the sexes. Thus, males formed gonopods in the first abdominal somite and lost the paired vestigial pleopods in somites 3–5 from the fourth instar; females retained the pleopods in somites 2–5, but these became biramous and had increased setation. The abdomen grew wider in females than in males from the fifth instar. Several morphological features of juveniles have phylogenetic and taxonomic implications: carapace motifs clearly place L. exaratus in the superfamily Xanthoidea, whereas the patterns of setation in the scaphognathite and first maxilliped epipod allow separation of this (xanthid) species from crabs of other Xanthoidean families.     

Diel activity patterns in Metapenaeus and Penaeus juveniles

Small (5–10.9 mm carapace length), medium (11–15.9 mm), and large (16–20.9 mm) juveniles of Metapenaeus anchistus, Metapenaeus sp., Penaeus monodon and P. merguiensis were stocked individually in glass tanks provided with sand substrate, sea water, artificial bamboo shelter, aeration and food. The seven activity types (recorded for each shrimp hourly for 24 h) were classified as below (burrowing) or above substrate (swimming, walking, stationary, in shelter, feeding and cleaning). Shrimp juveniles exhibited a strong diel periodicity — emergence and activity at night and burrowing in the day. The chi-square test showed that type of activity (above/below substrate) was associated with period (light/dark). Diurnal burrowing was greater among Metapenaeus than Penaeus; inversely, above substrate activities were more frequent for Penaeus species compared to Metapenaeus. Feeding was the major above substrate and nocturnal activity for M. anchistus, Metapenaeus sp. and P. monodon. Only P. Monodon used the shelter consistently. Frequency of the 7 activity types was dependent on juvenile size for Penaeus, e.g., the preference for shelters shifted to burrowing with increase in size in P. monodon. Results are discussed in relation to the importance of mangrove habitats in providing shelter to penaeids, in particular the mangrove-associated P. monodon and P. merguiensis.     

Four new species of Halicyclops Norman, 1903 (Copepoda,Cyclopoida) from Brazil

Four new species of Halicyclops are described and figured from plankton samples of the brackish waters of Pomonga River, Sergipe State (NE Brazil). The ornamentation of the median apical setae of the furcal rami is proposed as useful diagnostic character for species distinction.     

Masculinization of Female Crayfish,Procambarus Clarki (Girard)

Summary

Twenty-three immature female crayfish (Procambarus clarki (Girard); carapace length = 8.2–17.9 mm), implanted with androgenic gland implants obtained from P. clarki males, were observed for 326 days for evidence of masculinization. Number of molts completed ranged from three to eight. Of eleven surviving females, three were masculinized as evidenced by the partial transformation of their first pair of pleopods into male-type gonopods. No other evidence of masculinization, either internal or external was seen. Masculinization of the first pair of pleopods was unequal and first noted 290 days post-implantation. We conclude that androgenic gland implantation can induce masculinization of P. clarki females, albeit only with great difficulty. This is the first report of masculinization of female macruran decapods by androgenic gland implantation.     

Taxonomic analysis of species characters for copepodid instars 4 and 5 of the subfamily Eucyclopinae of European Russia

Morphological characters useful for taxonomic identification of older copepodid instars of the subfamily Eucyclopinae were studied among 14 species of Eucyclops, Macrocyclops, Ectocyclops, Paracyclops and Tropocyclops known from European Russia. For taxonomic analysis, the following elements of copepodid morphology were chosen: armament and proportion of furcal rami; morphology of swimming legs and reduced 5th and 6th legs; antennule segmentation; and relative body length of copepodid instars in comparison with the female length. Changes in morphology of major copepodid instars of the subfamily Eucyclopinae during ontogenetic development are traced and noticeable differences among five genera and 14 species are described. These differences among major copepodid stages may be important for both taxonomic and ecological analysis. For taxonomy, they provide information on development of sexes and species during maturation. For ecology they allow identification of the specimens at 4–5 copepodid instars in diapause. A key to major copepodid instars of the species from the subfamily Eucyclopinae which are known from limnetic habitats of the European part of Russia is presented.     

Das Schwimmen der Talitridae (Crustacea,Amphipoda): Funktionsmorphologie,Phänomenologie und Energetik

The Talitridae, well-known for their jumping behaviour, swim with help of the tail-flip. This movement of the abdomen is also known from other amphipods like the Gammaridae which are normally not able to move by jerks outside the water. The suspected homology between the tail-flip when swimming and the jerky movement of the abdomen when jumping gave rise to this investigation, mainly based on high frequency film recordings, on the swimming ofHyale nilssonii, Orchestia cavimana, andTalitrus saltator (family Talitridae) as well as three related species of the families Gammaridae and Corophiidae. Comparative morphometrical and SEM-studies on the habitus of the species and the build of the involved limbs reveal the rather uniform construction of the Gammaridea; functional adaptation to the environment and to the way of living become apparent in minor alterations. The joints of the pleopods and uropods show a clear structural adaptation to the mechanical strain during swimming. The pleopods are moved metachronally in all examined species; angular velocity and rate of beating indicate the efficiency of the swimming movement. In the Talitridae, the metachronal beat of the pleopods is nearly always coupled with the tail-flip while in the Gammaridae and Corophiidae the tail-flip, in addition to the beat of the pleopods, is mostly used for a start from the subsoil or for a change in swimming direction.H. nilssonii, Gammarus locusta, andCorophium volutator, all inhabitants of the tidal zone in the North Sea shallows, turned out to be the best swimmers while the (semi-) terrestrially living species,O. cavimana andT. saltator, proved to be rather poor swimmers. This clearly indicates the ecological significance of swimming for the different species. Furthermore, the tailflip is found to be of rather subordinate importance. It contributes to a higher velocity if used moderately but is rather obstructive if a large angle is covered while extending and flexing the abdomen. The efficiency of swimming is inversely proportional to the efficiency of jumping in the three talitridean species. Thus, better adaptation to terrestrial life is accompanied by loss of swimming efficiency. Examined under the aspect of locomotional homology, it is concluded that the tail-flip used while swimming is homologous to the jerky movement of the abdomen used for jumping. The comparison of the swimming performance of the examined species with other crustaceans and some fishes illustrates the over-all good results of the Gammaridea.     

Burrowing,Locomotion and Other Movements of the Echiuran Ochetostoma caudex

Burrowing, iocomotory and other movements of the echiuran Ochetostoma caudex have been examined and discussed. A continuous body cavity enables the worm to undergo peristaltic waves to pump water through the burrow without causing locomotion. The animal is capable of both forward and backward locomotion in its burrow. During forward locomotion, retrograde peristaltic waves are utilized which advance the animal in a step-wise fashion. Pressure changes within the coelom during burrowing, locomotion and during irrigation movements have been measured with the use of electronic recording techniques and the results interpreted in relation to direct visual observation. The structural and functional specializations for burrowing are discussed and compared with the activities of Priapulus caudatus, Sipunculus nudus and Bonellia viridis.     

Phyllocarida and the origin of the Malacostraca

Tentative homologies with Decapoda are proposed for grooves on the carapace of Echinocaris and Montecaris. Together with F.R. Schram's hoploid trend, reinforced by the discovery of a raptorial limb in Sairocaris, this may strengthen a phyllocarid origin for Eumalacostraca. Both these trends, however, may be parallelisms, and phylogenetic analysis is required. Pleopods are here proved to have been present in additional Archaeostraca, which improves their ancestral condition. E. Dahl's view of the phyllocarids as an evolutionary dead end is falsified from fossils, and the phyllocarids are supported as a stem group of the Malacostraca.     

Burrowing in frogs

More than 95% of burrowing Anura dig hindfeet first into the soil, a pattern unique to frogs among terrestrial vertebrates. The postero-laterally placed hindlimbs and associated musculature of frogs are preadaptations for hindfeet digging. One fossorial, backwards burrower, Glyphoglossus molossus (Microhylidae), has morphological modifications of the hindlimb for positioning the spade-like metatarsal tubercle and for increasing the force of the lower leg during digging. In contrast, in the headfirst burrower Hemisus marmoratus (Ranidae) there is extensive reorganization of the pectoral-cranial morphology compared to that: of a non-burrowing confamilial species. A model links the shifts in the pectoral morphology in Hemisus marmoratus to specific action patterns of headfirst: burrowing. Finally, data on stomach contents, natural history and energy utilization of frog species are presented to demonstrate the interrelationships of distinct loco. motor patterns with specific feeding strategies.     

Kinematics of swimming in two burrowing anguilliform fishes

Anguilliform or eel-like fishes are typically bottom dwellers, some of which are specialized burrowers. Although specializations for burrowing are predicted to affect the kinematics of swimming, it remains unknown to what extent this is actually the case. Here we examine swimming kinematics and efficiency of two burrowing anguilliform species, Pisodonophis boro and Heteroconger hassi, with different degrees of specialization for burrowing. Our data suggest that differences in the swimming kinematics may indeed be related to the differences in burrowing specialization and style between both species. The resemblance between the swimming kinematics of P. boro and previously published data for Anguilla anguilla and Anguilla rostrata may be linked with the relatively limited burrowing specialization of P. boro and suggests an overall stereotypy in anguilliform forward-swimming patterns. The body of H. hassi, in contrast, is more specialized for tail-first burrowing and backward swimming bears a striking resemblance to the backward burrowing motions observed in this species. These motions differ significantly from backward swimming in Anguilla and in P. boro. The kinematics of forward swimming are, however, comparable across species. Thus, our data suggest that specializations for burrowing may affect swimming kinematics in anguilliform fishes, but also that forward swimming and burrowing are not necessarily incompatible. Future studies comparing the kinematics and mechanics of burrowing in these and other anguilliform fishes are needed to better understand how specializations for burrowing constrain backward swimming in H. hassi.     

Pseudomoraria triglavensis gen. n., sp. n. (Copepoda,Harpacticoida) from a high-alpine reservoir in Slovenia

Numerous specimens of Pseudomoraria triglavensis gen.n., sp.n. were collected from small, high-alpine, reservoir Moilec. The reservoir is situated in the centre of Triglav national park (NW Slovenia) at an altitude of 1690 m. The new genus differs from related genera Moraria, Morariopsis and Paramorariopsis by the reduced articulation of exopodites and endopodites in both sexes. It also differs from Moraria by possessing sexually dimorphic furcal rami. Detail taxonomic differences between these related genera and their ecology are discussed.     

Species specific behavioural patterns (digging and swimming) and reaction to novel objects in wild type, Wistar, Sprague-Dawley and Brown Norway rats

Background

The purpose of the present study was to analyse species-specific forms of behaviour (digging and swimming) and response to novelty in laboratory rats and their wild type counterparts at a very early stage of laboratorization. Three behavioural phenomena were taken into account: burrowing, spontaneous swimming, and neophobic behaviour.

Principal Findings

Wild-type rats and three strains of laboratory rats were involved in experiments: Warsaw-Wild-Captive-Pisula-Stryjek (WWCPS), Wistar, Sprague-Dawley, and Brown Norway rats were compared in spontaneous swimming test, while WWCPS and Wistar rats were studied in burrowing and neophobia experiments. Wild rats were found to be faster at building tunnels than Wistar rats and at constructing more complex burrow systems. The experiment on neophobia showed that Wistar rats exhibited less neophobic responses and were more often trapped. WWCPS rats showed highly neophobic behaviour and were rarely trapped in this experiment. The experiment on swimming showed that WWCPS rats showed more complex water tank related activity than their laboratory counterparts. They swam and explored under surface environment.

Conclusions

The three experiments showed profound behavioural differences in quasi-natural forms of behaviour between wild type rats (WWCPS) and three laboratory strains frequently used in behavioural studies.     

A new species of Wellsopsyllus (Copepoda, Harpacticoida, Paramesochridae) from the deep Southern Ocean and remarks on its biogeography

The new deep-sea copepod species of the family Paramesochridae (Copepoda, Harpacticoida) presented here was collected during the scientific ANtarctic benthic DEEP-sea biodiversity cruise II (ANDEEP II) to the Weddell Abyssal Plain and western Weddell Sea and the CROZet natural iron bloom and EXport experiment (CROZEX cruise) to the Crozet Isles in the Indian Ocean. The new species has been allocated to the subgenus Wellsopsyllus within the genus Wellsopsyllus Kunz, 1981, based on the 1-segmented endopods and 3-segmented exopods of swimming legs 2–4. Furthermore, the new species shows a 1-segmented exopod of the antenna. Wellsopsyllus (W.) antarcticus sp. n. can be distinguished from its congeners by its small body size, the presence of two setae on the distal segment of the endopod of the first swimming leg, by the very small furcal rami, the extremely reduced endopods of second and third swimming legs, the reduced mandibular palp and maxillula, as well as by the size and armature of the fifth and sixth swimming legs in both sexes. Difficulties of the placement of the new species into a suitable genus show an insufficiency in the present family classification. In the future, a revision of the genus Wellsopsyllus is essential. With its presence in the western Weddell Sea and the Indian Ocean, the new species may have a wide distribution range. Furthermore, it is the first abyssal species of the genus Wellsopsyllus sampled outside of the Scotia Sea.     

Environmental factors affect swing gates as a barrier to large carnivores entering game farms

Burrowing animals such as warthog (Phacochoerus africanus), Cape porcupine (Hystrix africaeaustralis) and aardvark (Orycteropus afer) are able to compromise the integrity of fenced‐in farmlands by digging holes under game fences. These holes provide access for predators to enter the farm where they can kill livestock or captive game animals. Data collected from the use of swing gates (n = 263) installed along a 23.93 km game fence in the Otjozondjupa region of Namibia was analysed to determine the factors that influenced their efficacy at reducing hole creation along the fence by digging animals. Statistical analyses revealed that soil substrate, grass height, vegetation density, distance to the nearest permanent water source and season influenced digging activity along the fence line. The number of holes created and reopened decreased over time from the start of the study period, probably demonstrating that burrowing animals had learnt to use the swing gates rather than dig holes under the fence. These factors can inform the correct future usage of swing gates as a large predator exclusion method to ensure that they do not enter game farms, which will reduce the need to lethally control carnivores and burrowing animals.     

Comparison of the structure and presumptive function of Type II microtrichs and associated cuticular microstructures in species of <Emphasis Type="Italic">Gammarus</Emphasis> (Amphipoda,Crustacea) from different habitats

Type II microtrichs are slender, peg-like sensilla found on the exoskeleton of amphipod crustaceans. Using scanning electron microscopy, the microtrichs and associated cuticular microstructures were compared in six species and two subspecies of Gammarus known to live in different habitats, ranging from freshwater lentic and lotic to marine intertidal. In all the species, the microtrichs were clustered in arrays on the lateral and dorsal areas of the exoskeleton, and there were similarities in their configuration: semi-circular, L-shaped, and linear. Intraspecific variation of sensillum numbers and arrangement in arrays was slight compared with interspecific differences, where axial and lateral orientations of the microtrichs are believed to be related to current detection. Associated cuticular microstructures included in-common finely pixelated, polygonal plates, modified in some species by lateral stretching, the development of short spines, a scale-like appearance, and raised triangular peaks some of which may be related to locomotion, such as burrowing. Grouping of the eight species/subspecies along phylogenetic lines showed little support based on relatedness centred around calceolus-possession or water salinity requirements. A significant microstructural difference was observed between subspecies of Gammarus lacustris.