The Grapsidae and Ocypodidae (Decapoda: Brachyura) of Tanzania

Twenty-nine species of the Grapsidae and 15 species of the Ocypodidae were collected from the littoral zone in the area around Dar es Salaam. Nine species of Grapsidae and three of Ocypodidae were new records for the East African area, bringing the total of recorded species for the region to 35 and 26 respectively. The distribution of these crabs indicates a very limited endemism in the western Indian Ocean, with the majority of species forming an attenuated extension of the abundant Indo/West Pacific brachyuran fauna.
The habitat preference of each species is described. Within each subfamily there is a relative uniformity in general habitat preference, mode of life and manner of feeding. The species of each subfamily are then separated by substrate choice and preferred level on the shore. The greatest numbers of species occur in the most diverse habitats—the creek mangrove, the coastal mangrove and the reef flat.     

Male courtship cycles in three species of tropical Ilyoplax crabs (Decapoda,Brachyura, Ocypodidae)

The courtship behaviour and cycles of male courtship activity and colouration of Ilyoplax orientalis, I. delsmani and I. gangetica were studied in the field in Malaysia and Thailand. Each species had a distinctive chela waving or beckoning display. Depending on species, the chelipeds, carapaces, or both of waving males blanched to white in contrast to the cryptic colour of nonwaving males and females. All three of these tropical Ilyoplax exhibited semilunar cycles in male waving activity at the colony level. It was confirmed for I. orientalis that individual males cycled each semilunar period between waving and non-waving phases and exhibited different behaviour toward females during these two behavioural phases.     

Carapace movements aid air breathing in the semaphore crab,Heloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Summary While on land and recirculating branchial water the Australian semaphore crab Heloecius cordiformis (Decapoda: Ocypodidae), a semi-terrestrial airbreathing mangrove crab, sequentially depresses and elevates its carapace in a regular pump-like manner. The functional role of these carapace movements in aerial oxygen consumption is investigated. Carapace immobilisation (reversible and non-injurious) did not appear to affect branchial water circulation. In dry crabs (branchial water removed) carapace immobilisation had no effect on the rate of oxygen consumption (VO₂), heart rate or whole-body lactate (WBL) levels. In wet crabs (with branchial water) carapace immobilisation caused VO₂ to drop by 38% from 81 to 46 l O₂ · g^-1 · h^-1, heart rate to decline by 32%, from 2.5 to 1.7 Hz, and WBL levels to increase over 2.5-fold, from 0.27 to 0.67 mg · g^-1, after 3 h of carapace immobilisation. The (VO₂) of carapace-immobilised crabs with branchial water was similar to lung-occluded crabs with branchial water. Severe hypoxia induced physiological responses similar to those of carapace-immobilised crabs with branchial water. After 3 h of severe hypoxia, heart rate had declined by 80%, from 2.2 to 0.43 Hz, and the incidence of carapace pumping slowed by 85%, from 2.4 to 0.37 cycles · min^-1. It is concluded that in the absence of carapace movements branchial water in some way inteferes with lung ventilation. Under normal circumstances water circulation and lung ventilation are mutually exclusive processes (due to their singular dependence on the scaphognathites), yet in Heloecius these processes must be carried out simultaneously. Carapace movements may alleviate this conflict.Abbreviations FF, FR, SF, SR fast-forward, fast-reverse, slow-forward, slow-reverse scaphognathite pumping - MEA Milne Edwards aperture - VO₂ rate of oxygen consumption - WBL whole-body lactate     

Substratum preference of the tropical estuarine crabs,Uca tangeri Eydoux (Ocypodidae) and Ocypode cursor Linne (Ocypodidae)

The vertical and horizontal distribution of two burrowing mud crabs, (Ocypodidae) and Ocypode cursor Linne 1758 (Ocypodidae) are described for the Bonny Estuary (7° 00' E: 4° 20′ N), S. Nigeria. Substratum preference is the most prominent factor influencing distribution, but lack of tolerance to low salinities (< 6%_o) is also important, and prevents Ocypode cursor from occurring close to freshwater. The two species have slightly different sediment organic content and grade size requirements. Ocypode cursor was concentrated in well drained sandy sediment above mid-tide-level, whilst Uca tangeri was found in water-logged areas slightly above and below Mid-tide-level. This vertical demarcation is attributed to differential feeding and burrowing adaptations related to different substrata, rather than to differential tolerances to desiccation.     

Histologische Untersuchungen zum Bau des Auges von Ocypode cursor (Brachyura)

Zusammenfassung Im Ommatidium des Komplexauges von Ocypode cursor wurde entgegen einer früheren Untersuchung eine achte Retinulazelle gefunden. Sie unterscheidet sich nach Form und Lage von den sieben regulären Retinulazellen. Ihr Kontakt zum Rhabdom und der Besitz eines Axons (bisher bei Decapoden unbekannt) widerlegen die für andere Decapoden geäußerte Ansicht, daß diese Zelle rudimentär sei. Ihre besondere Ausbildung legt den Gedanken nahe, daß sie funktionell spezialisiert ist.Am distalen Ende der Retinula liegt eine im Schnitt quer zur Ommatidienachse kreuzförmige pigmentfreie Zone (Abb. 3, 4). Ein Arm, in jedem Ommatidium der gleiche, enthält einen kugeligen Kern. Es ist anzunehmen, daß diese pigmentfreie Zone den Zellkörper der achten Retinulazelle darstellt. Diese Zelle steht in räumlichem Kontakt zum Rhabdom. Der Raum zwischen den vier Armen der Zelle 8 wird von den distalen, stark pigmentierten Enden der sieben regulären Sinneszellen gefüllt. Die in der Regel größere Zelle 7 liegt hinten, Zellen 1 und 2 oben, Zellen 3 und 4 vorn, Zellen 5 und 6 unten. Die Kerne dieser Zellen sind länglich ellipsoid. Der kernhaltige Arm der Zelle 8 verjüngt sich nach proximal zu einem Axon (Abb. 6), das an der Peripherie des Rings der sieben regulären Retinulazellen an der Naht zwischen den Zellen 6 und 7 zur Basalmembran zieht (Abb. 7). Unmittelbar über der Basalmembran divergieren die Retinulazellen: Zelle 1 zieht nach hinten oben, Zellen 2 und 3 nach vorn oben, Zellen 4 und 5 nach vorn unten, Zellen 6 bis 8 nach hinten unten (Abb. 9). In dieser asymmetrischen Gruppierung durchstoßen die Axone der Retinulazellen die vier lanzettlichen Öffnungen, die zur Basis der in der darüber und darunter liegenden Reihe nächst benachbarten Ommatidien ziehen (Abb. 10). Auf diese Weise ergibt sich ein regelmäßiger Wechsel von Öffnungen in der Basalmembran mit drei und fünf Querschnitten von Axonen (Abb. 11).

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Summary In the ommatidium of the apposition eye of Ocypode cursor eight retinula cells are found, where the eighth accessory cell has a characteristic shape and position distinct from the other seven retinula cells. For other decapods this cell has been assumed to be rudimentary. At least for Ocypode its contact with the rhabdome and the possession of an axon contradict this assumption. A functional specialization of this cell seems more probable.In sections perpendicular to the optical axis a cross-shaped pigmentless structure appears at the distal end of the retinula. One bar of this cross (in any particular ommatidium the same) contains a spherical nucleus (Figs. 3, 4). All four bars seem to be in contact with the central rhabdome. The similar appearance of all four bars and the presence of only one nucleus in this region favour the assumption that all four bars belong to one and the same cell. The space between the bars is filled by the densely pigmented distal ends of the seven regular retinula cells: the dorsal space by cells 1 and 2, the anterior by 3 and 4, the ventral by 5 and 6, and the posterior by cell 7. The nuclei of these cells are elongated. The nucleus-containing bar tapers proximally into an axon (Fig. 6) which extends towards the basilar membrane peripheral to the rosette of the seven regular sense cells, close to the border of cells 6 and 7 (Fig. 7). Immediately adjacent to the basement membrane the retinula cells (in this case, their axons) diverge peripherally: the direction of No. 1 becomes dorso-posterior, No. 2 and No. 3 dorso-anterior, No. 4 and 5 ventro-anterior, No. 6–8 ventro-posterior (Fig. 9). In this asymmetric distribution the axons of the retinular cells pass through the basement membrane by openings which extend between the bases of neighbouring ommatidia in the next higher or lower row of ommatidia (Fig. 10). Thus a sequence of openings follows with alternately three and five cross sections of axons (Fig. 11).

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Herrn E. Freiberg danke ich für die Ausführung der Zeichnungen, Frl. I. Geiss und Drs. R. und S. Pickering für Hilfe bei der Anfertigung des Manuskriptes, Herrn Dr. K. Kirschfeld für dessen kritische Durchsicht.     

Beobachtungen zur Biologie und zum Verhalten der Klippenkrabbe Grapsus grapsus L. (Brachyura Grapsidae) auf Galapagos und am ekuadorianischen Festland1)2

• 1 Während eines einjährigen Aufenthaltes auf den Galapagosinseln wurde die Klippenkrabbe Grapsus grapsus L. unregelmäßig und am ekuadorianischen Festland bei Palmar in der Nähe von Guayaquil fünf Tage lang beobachtet.
• 2 G. grapsus bewohnt die Gezeitenzone von Felsküsten und lebt im wesentlichen von dem hier gedeihenden Algenbewuchs.
• 3 Landlebende Feinde sind auf Galapagos vor allem die beiden kleinen Reiher Butorides sundevalli und Nyctanassa violacea. Im Wasser drohen Raubfische (z. B. Cirrhitus rivulatus) und Kraken. Auf dem ekuadorianischen Festland kommen Kleinbären als Landfeinde vor.
• 4 Die Galapagos-Klippenkrabben sind, verglichen mit den ekuadorianischen Artgenossen, vor allem prächtiger gefärbt und bedeutend größer. Zum Teil variieren sie farblich ein wenig von Insel zu Insel.
• 5 Als bemerkenswerte sekundäre Geschlechtsmerkmale werden Größen-unterschiede und dichte, an den Propoditen der ersten beiden Laufbeinpaare befindliche Bürsten beschrieben, die den ausgewachsenen ♀♀ fehlen.
• 6 An den Oberseiten der Laufbeine stehen Reihen von Haaren, die wahrscheinlich der Wahrnehmung des gegenseitigen Betastens mit den Daktylopoditen dienen.
• 7 Klippenkrabben gehen langsam vorwärts oder etwas schräg, auf der Flucht oder als Verfolger rennen sie seitwärts. Sie springen geschickt von Fels zu Pels und schwimmen mit raschen Schlägen der dorsoventral abgeflachten Laufbeine kürzere Strecken über die Wasseroberfläche hinweg.
• 8 Die Krabben zupfen mit den vorn beißzangenartig verbreiterten Scheren Algenbewuchs ab. Sie fressen gerne Fleisch und verfolgen und ergreifen alles, was klein ist und sich in ihrer Nähe bewegt. Insbesondere die großen ♂♂ überfallen kleinere Artgenossen und fressen sie ganz oder deren autotomierte Beine.
• 9 Vor allem kleine und mittelgroße Tiere halten sich bevorzugt in einer Gruppe mit gleich großen Artgenossen auf und sind meistens alle gleich orientiert.
• 10 In vielfältigen Situationen berühren die Krabben einander mit den Laufbeinen und erteilen dabei anscheinend mechanische und eventuell chemische Signale. Eine kleine Krabbe kann wohl durch seitliches Betasten einen drohenden Angriff eines großen Artgenossen abwenden.
• 11 Kraftproben zwischen ♂♂ werden meist durch einen Kommentkampf entschieden, in dem drei deutlich unterscheidbare Ausdrucksbewegungen auftreten können.
• 12 Bei ungenügender optischer Kontrolle der Umgebung oder auf der Flucht zeigen die Krabben spezielle Abwehrbewegungen.
• 13 In der ersten Phase der Balz verfolgt das imponierende ♂ das ♀, in der zweiten folgt das ♂ dem langsam zurückweichenden, weiter imponierenden ♂ und betastet es. Dann kann es zur Kopulation kommen. Kleine ♂♂ versuchen zuweilen, mit gleich großen und größren ♀♀ zu kopulieren, ohne daß ein Balzspiel vorangegangen ist.
• 14 In bestimmten Situationen spritzen die Krabben Wasserfontänen aus kleinen beweglichen Düsen gezielt auf Artgenossen oder auch spontan geradeaus von sich weg; die Düsen sitzen an den Basen der zweiten Antennen.
• 15 Beim normalen langsamen Gehen führt G. grapsus oft die leere Schere vom Boden zum Mund. Das ♂ zeigt solches ?Scheinfressen” auch bei der Kopula.
• 16 Ruhig sitzende Tiere sieht man zuweilen die Laufbeine aneinander reiben. Zusätzlich hüllen sie manchmal die Unterseite ihres Körpers und die Peraeopoden in Schaum ein, der aus den frontalen Öffnungen der Kiemenhöhlen tritt. Beides dürften Reinigungshandlungen sein.

     

Notes on the behaviour of Ocypode ryderi Kingsley (Crustacea,Brachyura)

The main aggressive patterns of Ocypode ryderi are described: the claws can be pointed downwards in a shield‐like position against a facing opponent or out stretched against an opponent coming from behind. While the first pattern often induces an analogous reaction in the opponent and can easily be followed by a fight consisting in reciprocal claws hitting, the latter always leads to avoidance of a further attack. A waving display occurs in courtship. The main defensive patterns against predators or men consists of a zig‐zag flight towards the sea or the burrow and a “terrifying” sound emitted by striking the claws together. Running and cleaning are executed as in the majority of the Ocypode. An attempt to compare these patterns with those of other species of Ocypode has been made.     

Earthen Structures Built by Ilyoplax dentimerosa (Crustacea,Brachyura, Ocypodidae)

Under observation, the small ocypodid crab Ilyoplax dentimerosa was found to commonly build three types of earthen structures: a barricade near its neighbour's burrow, a fence at an intermediate position between the burrows of the builder and its neighbour, and a minishelter near the builder's burrow. The sex ratio of barricade builders was found to be close to 1:1, whereas most of the fence builders were found to be female. Crabs against which barricades and fences were built, were usually smaller than the builders. Both barricade builders and fence builders had, in most cases, minishelters at the side, facing the barricade or the fence. Removal and rebuilding experiments for barricades and fences demonstrated that both structures functioned to deter the approach of the builder's neighbour to the builder's activity site. Fences also had the effect of restraining the movement of the builder toward the fence site.     

Relative growth before and after sexual maturity in Ocypode platytarsis (Milne Edwards) and Ocypode cordimana (Desmarest) (Crustacea: Decapoda)

The relative growth of two species of the genus Ocypode inhabiting different ecological niches was studied before and after sexual maturity. The growth coefficient of both sexes of O. cordimana, which inhabits supra-littoral zones, is higher than that of O. platytarsis found in wave-wash zones.s

Some characteristics such as eyestalk length, major chela width and obdomen width do not show any difference in the growth pattern of either species of Ocypode at the onset of sexual maturity. Other characteristics such as minor and major chela length in male O. platytarsis; carapace length, minor chela length and width in female O. platytarsis; major chela length and third walking leg length in male O. cordimana showed a difference in the growth pattern after the onset of sexual maturity.

Major chela length of males of both species is clearly a sexually dimorphic feature since it showed an increase in the growth coefficient after sexual maturity. Merus length in male O. platytarsis and minor chela length in female O. cordimana also showed such sexual dimorphism.

There are several characteristics in which the growth coefficient declines after sexual maturity. The change in growth coefficient of different characteristics may depend on the adaptive value of the characteristic in reproductive or post-reproductive activities of O. cordimana and O. platytarsis.     

Carapace movements associated with ventilation and irrigation of the branchial chambers in the semaphore crab,Heloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Summary Carapace movements in crabs are briefly reviewed. While on land and recirculating branchial water, the Australian semaphore crab Heloecius cordiformis (Decapoda: Ocypodidae), a semi-terrestrial air-breathing mangrove crab, sequentially depresses and elevates its carapace relative to its thorax (0.5–1 mm excursion) in a regular pump-like manner. In quiescent crabs each carapace-pumping cycle lasts about 4 s; carapace depression takes 3 s and elevation 1 s. Carapace movements are brought about by pressures generated within the branchial chambers by the scaphognathites, probably in combination with carapace muscles. Carapace movements are associated with bilaterally synchronised scaphognathite activity. Unilateral scaphognathite activity was not observed. During normal forward recirculation of branchial water the scaphognathites beat at about 1.5 Hz (slow-forward pumping) and the lungs (epibranchial chambers) are not ventilated. In Heloecius, the lungs are not physically separated from the gills below by an anatomical barrier. Lung ventilation is accomplished during the following sequence of events: the carapace is lowered and the scaphognathites pump in a fast-forward mode at about 2.8 Hz. This activity preferentially pumps air out of the lungs and generates suction within the branchial chambers (4–10 cm H₂O below ambient) which draws water from external body surfaces into the hypobranchial space below and around the gills. At the end of the carapace's downward travel the scaphognathites switch from fast-forward to fastreverse beating at about 4 Hz. This pumps air into the lungs and the carapace elevates. As a result, during carapace elevation the water which had previously been drawn into the branchial chambers by fast-forward pumping activity is released and flows out between the legs and into the abdominosternal cavity. When the carapace reaches its original resting or up position the scaphognathites switch from fast-reverse to slowforward beating to re-establish water recirculation through the branchial chambers. This cycle is subsequently repeated. In stationary crabs, there are 2 carapace-pumping cycles per minute, increasing to 14 per minute in active crabs (walking). When water is absent, the lungs are preferentially ventilated by slow-reverse scaphognathite pumping activity. Carapace movements do not occur in the absence of branchial water. Carapace pumping is thought to provide a mechanism which permits the scaphognathites to ventilate the lungs in the presence of recirculating branchial water, without this water interfering with lung ventilation or being lost to the environment.Abbreviations FF, FR, SF, SR fast-forward, fast-reverse, slowforward, slow-reverse scaphognathite pumping - MEA Milne Edwards aperture     

Polymorphic microsatellite DNA markers in the mangrove crab Ucides cordatus (Brachyura: Ocypodidae)

The isolation and characterization of the first polymorphic microsatellite markers for the mangrove crab Ucides cordatus are described. The number of alleles at each locus ranged from three to 25, mean of nine alleles, in 46 crabs captured in two Brazilian mangroves. The markers averaged high levels of observed (0.709 ± 0.183) and expected (0.716 ± 0.170) heterozygosities. Departures from Hardy-Weinberg equilibrium were observed at two loci. Linkage disequilibrium tests were not significant and no evidence of null alleles was detected. All these microsatellite loci are expected to be useful in estimating fine-scale population processes of this valuable mangrove species currently subjected to excessive fishing efforts.     

Vertical distribution by demographic groups of ghost crab Ocypode quadrata (Crustacea: Brachyura)

The ghost crab Ocypode quadrata plays an important role in energy transfer between trophic levels, and has been widely used in evaluations of impacted environments. In order to provide data on the biology of this potential bioindicator species, the population structure and vertical distribution of individuals were studied on two beaches in southeastern Brazil. Each beach was divided into quadrants of 1000 m² with boundaries of upper, middle and lower levels in relation to the waterline. Collected monthly by active searching through one year, the specimens of O. quadrata were sexed, measured for carapace width, and returned to the beach. Of the total of 1904 specimens collected, the largest proportion (46.2%) were males, followed by 31.4% juveniles. The vertical distribution of the ghost crabs differed among age groups: males mostly occupied the middle and upper levels; adult females, ovigerous or not, were more abundant in the lower level; and juveniles were evenly distributed in all levels, with a slight tendency toward the middle. The sex ratio favored males in a few months of the year and in the larger size classes. The abundance of O. quadrata is limited by low temperatures, and its spatial and temporal distribution is controlled by food availability and ease of reproduction. Knowledge of the biology of these crabs is essential in order to use them as a bioindicator species; the vertical distribution patterns may reflect changes in the beach hydrodynamics or other environmental factors.     

Spermatogenesis of the spider crab Maja brachydactyla (Decapoda: Brachyura)

This study describes spermatogenesis in a majid crab (Maja brachydactyla) using electron microscopy and reports the origin of the different organelles present in the spermatozoa. Spermatogenesis in M. brachydactyla follows the general pattern observed in other brachyuran species but with several peculiarities. Annulate lamellae have been reported in brachyuran spermatogenesis during the diplotene stage of first spermatocytes, the early and mid‐spermatids. Unlike previous observations, a Golgi complex has been found in mid‐spermatids and is involved in the development of the acrosome. The Golgi complex produces two types of vesicles: light vesicles and electron‐dense vesicles. The light vesicles merge into the cytoplasm, giving rise to the proacrosomal vesicle. The electron‐dense vesicles are implicated in the formation of an electron‐dense granule, which later merges with the proacrosomal vesicle. In the late spermatid, the endoplasmic reticulum and the Golgi complex degenerate and form the structures–organelles complex found in the spermatozoa. At the end of spermatogenesis, the materials in the proacrosomal vesicle aggregate in a two‐step process, forming the characteristic concentric three‐layered structure of the spermatozoon acrosome. The newly formed spermatozoa from testis show the typical brachyuran morphology. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

The feeding behaviour of the sand crab Scopimera inflata (Decapoda, Ocypodidae)

A sequence of activities all related to the feeding of Scopimera inflata H. Milne-Edwards, 1873 can be distinguished during the daytime periods of low tide. Emergence begins one to two hours after a feeding area is uncovered, reaches a peak after a further three hours but some crabs are still emerging after the tide has turned. Before feeding commences the burrow is cleared of waste sand down to the water-table. While feeding S. inflata forms a feeding trench which acts as an escape route back to the hole and is defended against other crabs of the same species. Agonistic behaviour becomes evident as feeding progresses. Threat displays, fighting, displacement feeding, and agitation feeding are recognizable behaviour patterns which have been observed.     

Genetic relationship among Japanese sentinel crabs (Decapoda: Ocypodidae: genus Macrophthalmus)

Seven species (eight populations) of sentinel crabs (genus Macrophthalmus) from the Japan coast and Uca vocans and Ocypode ceratophthalma, were examined electrophoretically for genetic variations in 13 enzymatic and one non-enzymatic protein comprising 17 loci. Most species were highly differentiated from each other (Nei's genetic distance, 0.29-1.63). The least genetic distance was found between M. japonicus and M. banzai, the genetic distinctiveness of the two taxa being supported by three divergent loci with no common allele. The genetic relationships among Macrophthalmus species differed greatly from those inferred from morphological features, with a UPGMA tree suggesting that the sub-genus Macrophthalmus is polyphyletic.     

Sperm ultrastructure of the spider crab Maja brachydactyla (Decapoda: Brachyura)

This study describes the morphology of the sperm cell of Maja brachydactyla, with emphasis on localizing actin and tubulin. The spermatozoon of M. brachydactyla is similar in appearance and organization to other brachyuran spermatozoa. The spermatozoon is a globular cell composed of a central acrosome, which is surrounded by a thin layer of cytoplasm and a cup‐shaped nucleus with four radiating lateral arms. The acrosome is a subspheroidal vesicle composed of three concentric zones surrounded by a capsule. The acrosome is apically covered by an operculum. The perforatorium penetrates the center of the acrosome and has granular material partially composed of actin. The cytoplasm contains one centriole in the subacrosomal region. A cytoplasmic ring encircles the acrosome in the subapical region of the cell and contains the structures‐organelles complex (SO‐complex), which is composed of a membrane system, mitochondria with few cristae, and microtubules. In the nucleus, slightly condensed chromatin extends along the lateral arms, in which no microtubules have been observed. Chromatin fibers aggregate in certain areas and are often associated with the SO‐complex. During the acrosomal reaction, the acrosome could provide support for the penetration of the sperm nucleus, the SO‐complex could serve as an anchor point for chromatin, and the lateral arms could play an important role triggering the acrosomal reaction, while slightly decondensed chromatin may be necessary for the deformation of the nucleus. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.