Lower Ordovician synziphosurine reveals early euchelicerate diversity and evolution.

Euchelicerata is a clade of arthropods comprising horseshoe crabs, scorpions, spiders, mites and ticks, as well as the extinct eurypterids (sea scorpions) and chasmataspidids. The understanding of the ground plans and relationships between these crown-group euchelicerates has benefited from the discovery of numerous fossils. However, little is known regarding the origin and early evolution of the euchelicerate body plan because the relationships between their Cambrian sister taxa and synziphosurines, a group of Silurian to Carboniferous stem euchelicerates with chelicerae and an unfused opisthosoma, remain poorly understood owing to the scarce fossil record of appendages. Here we describe a synziphosurine from the Lower Ordovician (ca. 478 Ma) Fezouata Shale of Morocco. This species possesses five biramous appendages with stenopodous exopods bearing setae in the prosoma and a fully expressed first tergite in the opisthosoma illuminating the ancestral anatomy of the group. Phylogenetic analyses recover this fossil as a member of the stem euchelicerate family Offacolidae, which is characterized by biramous prosomal appendages. Moreover, it also shares anatomical features with the Cambrian euarthropod Habelia optata, filling the anatomical gap between euchelicerates and Cambrian stem taxa, while also contributing to our understanding of the evolution of euchelicerate uniramous prosomal appendages and tagmosis.

An earliest Triassic age for Tasmaniolimulus and comments on synchrotron tomography of Gondwanan horseshoe crabs.

Constraining the timing of morphological innovations within xiphosurid evolution is central for understanding when and how such a long-lived group exploited vacant ecological niches over the majority of the Phanerozoic. To expand the knowledge on the evolution of select xiphosurid forms, we reconsider the four Australian taxa: Austrolimulus fletcheri, Dubbolimulus peetae, Tasmaniolimulus patersoni, and Victalimulus mcqueeni. In revisiting these taxa, we determine that, contrary to previous suggestion, T. patersoni arose after the Permian and the origin of over-developed genal spine structures within Austrolimulidae is exclusive to the Triassic. To increase the availability of morphological data pertaining to these unique forms, we also examined the holotypes of the four xiphosurids using synchrotron radiation X-ray tomography (SRXT). Such non-destructive, in situ imaging of palaeontological specimens can aid in the identification of novel morphological data by obviating the need for potentially extensive preparation of fossils from the surrounding rock matrix. This is particularly important for rare and/or delicate holotypes. Here, SRXT was used to emphasize A. fletcheri and T. patersoni cardiac lobe morphologies and illustrate aspects of the V. mcqueeni thoracetronic doublure, appendage impressions, and moveable spine notches. Unfortunately, the strongly compacted D. peetae precluded the identification of any internal structures, but appendage impressions were observed. The application of computational fluid dynamics to high-resolution 3D reconstructions are proposed to understand the hydrodynamic properties of divergent genal spine morphologies of austrolimulid xiphosurids.

Histological characterization of the gastrointestinal tract of the adult horseshoe crab (Limulus polyphemus) with special reference to the stomach.

The American horseshoe crab (Limulus polyphemus) is one of four extant species in the Order Xiphosura, subphylum Chelicerata, and are evolutionarily more closely related to scorpions and spiders, than crabs. The basic structure, function, and physiology of these invertebrates and their internal organs are not well documented in the literature. In this study, the gastrointestinal system, with a focus on the stomach, of adult L. polyphemus were assessed by gross and histologic methods to further characterize the pyloric valve, the lining of the ventricular lumen, and the muscular tunics of the stomach. Determination of normal anatomical structure of this organ system, along with characterization of the esophagus and intestinal tract, will set a standard against which tissue abnormalities, such as those seen with disease or pathology were to arise, would allow for better interpretation.

Environmental factors and occurrence of horseshoe crabs in the northcentral Gulf of Mexico.

This study provides regional-scale data on drivers of horseshoe crab (Limulus polyphemus) presence along the northcentral Gulf of Mexico coast and has implications for understanding habitat suitability for sparse horseshoe crab populations of conservation concern worldwide. To collect baseline data on the relationship between environmental factors and presence of horseshoe crabs, we surveyed four sites from the Fort Morgan peninsula of Mobile Bay, Alabama (AL) to Horn Island, Mississippi (MS). We documented number, size and sex of live animals, molts, and carcasses as metrics of horseshoe crab presence and demographics for two years. Data were compared to in situ and remotely sensed environmental attributes to assess environmental drivers of occurrence during the time of study. Overall, greater evidence of horseshoe crab presence was found at western sites (Petit Bois and Horn Islands) compared to eastern sites (Dauphin Island, Fort Morgan peninsula), mediated by a combination of distance from areas of high freshwater discharge and interannual variation in weather. Higher sex ratios also were found associated with higher occurrence, west of Mobile Bay. Land cover, particularly Bare Land and Estuarine Emergent Wetland classes that are common to western sites, was most predictive of live animal and to some extent carcass occurrence. Our findings suggest that small-scale variation in habitat quality can affect occurrence of horseshoe crabs in sparse populations where density is not a limiting factor. Data from molts and carcasses were informative to supplement live animal data and may be useful to enhance ecological assessment and support conservation and management in regions with sparse populations.

A reappraisal of Paleozoic horseshoe crabs from Russia and Ukraine.

Xiphosura are extant marine chelicerates that have displayed apparent morphological conservatism and remarkable survivorship across their ~ 480 Ma fossil record. The easily recognisable features that are known to even the earliest xiphosurans-a crescentic prosoma and often trapezoidal thoracetron (opisthosoma)-have generated debate surrounding their origins and taxonomic significance. This interest resulted in the description of numerous horseshoe crab species during the early to mid-twentieth century, particularly in Russia, that have remained unrevised since their original publications and unconsidered in the light of recent phylogenetic hypotheses. Here, we re-examine the non-belinurid taxa housed within the Chernyshev Central Museum for Geological Exploration in Saint Petersburg. We present the first formal diagnosis of Bellinuroopsis rossicus, erect Shpineviolimulus jakovlevi (Glushenko and Ivanov, 1961) comb. nov., to contain the species formerly described as 'Paleolimulus' jakovlevi and refer Paleolimulus juresanensis to Paleolimulidae incertae sedis. Phylogenetic analysis places S. jakovlevi at the base of Limulina. This position, coupled with a prosomal shield that is notably larger than the thoracetron, and lack of hypertrophied genal spines, suggests that this morphology may represent the ancestral austrolimulid shape. As an extension of this revision, we assessed the general austrolimulid morphological characters and uncovered two possible groups of these bizarre xiphosurids.

Air Breathing in an Exceptionally Preserved 340-Million-Year-Old Sea Scorpion.

Arachnids are the second most successful terrestrial animal group after insects [1] and were one of the first arthropod clades to successfully invade land [2]. Fossil evidence for this transition is limited, with the majority of arachnid clades first appearing in the terrestrial fossil record. Furthermore, molecular clock dating has suggested a Cambrian-Ordovician terrestrialization event for arachnids [3], some 60 Ma before their first fossils in the Silurian, although these estimates assume that arachnids evolved from a fully aquatic ancestor. Eurypterids, the sister clade to terrestrial arachnids [4-6], are known to have undergone major macroecological shifts in transitioning from marine to freshwater environments during the Devonian [7, 8]. Discoveries of apparently subaerial eurypterid trackways [9, 10] have led to the suggestion that eurypterids were even able to venture on land and possibly breathe air [11]. However, modern horseshoe crabs undertake amphibious excursions onto land to reproduce [12], rendering trace fossil evidence alone inconclusive. Here, we present details of the respiratory organs of Adelophthalmus pyrrhae sp. nov. from the Carboniferous of Montagne Noire, France [13], revealed through micro computed tomography (µ-CT) imaging. Pillar-like trabeculae on the dorsal surface of each gill lamella indicate eurypterids were capable of subaerial breathing, suggesting that book gills are the direct precursors to book lungs while vascular ancillary respiratory structures known as Kiemenplatten represent novel air-breathing structures. The discovery of air-breathing structures in eurypterids indicates that characters permitting terrestrialization accrued in the arachnid stem lineage and suggests the Cambrian-Ordovician ancestor of arachnids would also have been semi-terrestrial.

Untangling the Gordian knot-further resolving the super-species complex of 300-million-year-old xiphosurids by reconstructing their ontogeny.

The group Xiphosurida (horseshoe "crabs") is today only represented by four species. However, in the fossil record, several dozen species have been described, especially from the Carboniferous (about 300 million years ago). Several species have been interpreted as representatives of Euproops or Belinurus, but there is ongoing discussion which of these species are valid and how they can be differentiated. Recent studies suggested that differences in the timing of individual development could provide information for species distinction, exemplified by studies on Euproops danae (Mazon Creek, USA) and Euproops sp. ("Piesproops"; Piesberg, Germany). For this study, we reinvestigated all Carboniferous xiphosurids from the British Coal Measures stored in the collections of the Natural History Museum London. Size comparisons of the specimens revealed nine size groups; the smaller specimens were originally labelled as Belinurus, the larger ones as Euproops. The nine size groups exhibit five different morphotypes differing in structures surrounding the posterior shield (= thoracetron): spines of different lengths and, in larger specimens, a more or less developed flange. Two of these morphotypes show significantly longer spines than the remaining specimens and could be conspecific as E. anthrax. The remaining specimens are interpreted as growth series of another species, presumably of E. rotundatus. An ontogenetic flange formation is also known from E. danae and the "Piesproops", but the timing differs between all three species. In E. rotundatus, the flange develops rather late, but then comparably abruptly, which makes this development more metamorphic in relation to development in the other species.

Xiphosurid from the Tournaisian (Carboniferous) of Scotland confirms deep origin of Limuloidea.

Horseshoe crabs are archetypal marine chelicerates with a fossil record extending from the Lower Ordovician to today. The major horseshoe crab groups are thought to have arisen in the middle to late Palaeozoic. Here we present the oldest known limuloid from the lower Carboniferous (Tournaisian stage, c. 350 million years ago) of Scotland: Albalimulus bottoni gen. et sp. nov. A comprehensive phylogenetic analysis supports the placement of A. bottoni as a representative of the extant family Limulidae and 100 million years older than any other limulid taxon. The use of geometric morphometric analyses corroborate the erection of the new taxon and illustrates the exploitation of morphospace by xiphosurids. This new taxon highlights the complex evolutionary history of xiphosurids and the importance of documenting these unique Palaeozoic individuals.

Insights into the 400 million-year-old eyes of giant sea scorpions (Eurypterida) suggest the structure of Palaeozoic compound eyes.

Sea scorpions (Eurypterida, Chelicerata) of the Lower Devonian (~400 Mya) lived as large, aquatic predators. The structure of modern chelicerate eyes is very different from that of mandibulate compound eyes [Mandibulata: Crustacea and Tracheata (Hexapoda, such as insects, and Myriapoda)]. Here we show that the visual system of Lower Devonian (~400 Mya) eurypterids closely matches that of xiphosurans (Xiphosura, Chelicerata). Modern representatives of this group, the horseshoe crabs (Limulidae), have cuticular lens cylinders and usually also an eccentric cell in their sensory apparatus. This strongly suggests that the xiphosuran/eurypterid compound eye is a plesiomorphic structure with respect to the Chelicerata, and probably ancestral to that of Euchelicerata, including Eurypterida, Arachnida and Xiphosura. This is supported by the fact that some Palaeozoic scorpions also possessed compound eyes similar to those of eurypterids. Accordingly, edge enhancement (lateral inhibition), organised by the eccentric cell, most useful in scattered light-conditions, may be a very old mechanism, while the single-lens system of arachnids is possibly an adaptation to a terrestrial life-style.

Lift and Drag Acting on the Shell of the American Horseshoe Crab (Limulus polyphemus).

The intertidal zone is a turbulent landscape where organisms face numerous mechanical challenges from powerful waves. A model for understanding the solutions to these physical problems, the American horseshoe crab (Limulus polyphemus), is a marine arthropod that mates in the intertidal zone, where it must contend with strong ambient flows to maintain its orientation during locomotion and reproduction. Possible strategies to maintain position include either negative lift generation or the minimization of positive lift in flow. To quantify flow over the shell and the forces generated, we laser-scanned the 3D shape of a horseshoe crab, and the resulting digital reconstruction was used to 3D-print a physical model. We then recorded the movement of tracking particles around the shell model with high-speed video and analyzed the time-lapse series using particle image velocimetry (PIV). The velocity vector fields from PIV were used to validate numerical simulations performed with the immersed boundary (IB) method. IB simulations allowed us to resolve the forces acting on the shell, as well as the local three-dimensional flow velocities and pressures. Both IB simulations and PIV analysis of vorticity and velocity at a flow speed of 13 cm/s show negative lift for negative and zero angles of attack, and positive lift for positive angles of attack in a free-stream environment. In shear flow simulations, we found near-zero lift for all orientations tested. Because horseshoe crabs are likely to be found primarily at near-zero angles of attack, we suggest that this negative lift helps maintain the orientation of the crab during locomotion and mating. This study provides a preliminary foundation for assessing the relationship between documented morphological variation and potential environmental variation for distinct populations of horseshoe crabs along the Atlantic Coast. It also motivates future studies which could consider the stability of the horseshoe crab in unsteady, oscillating flows.

On the appendicular anatomy of the xiphosurid Tachypleus syriacus and the evolution of fossil horseshoe crab appendages.

Xiphosurida-crown group horseshoe crabs-are a group of morphologically conservative marine chelicerates (at least since the Jurassic). They represent an idealised example of evolutionary stasis. Unfortunately, body fossils of horseshoe crabs seldom preserve appendages and their associated features; thus, an important aspect of their morphology is absent in explorations of their conservative Bauplan. As such, fossil horseshoe crab appendages are rarely considered within a comparative framework: previous comparisons have focussed almost exclusively on extant taxa to the exclusion of extinct taxa. Here, we examine eight specimens of the xiphosurid Tachypleus syriacus (Woodward, 1879) from the Cenomanian (ca 100 Ma) Konservat-Lagerstätten of Lebanon, five of which preserve the cephalothoracic and thoracetronic appendages in exceptional detail. Comparing these appendages of T. syriacus with other fossil xiphosurids highlights the conserved nature of appendage construction across Xiphosurida, including examples of Austrolimulidae, Paleolimulidae, and Limulidae. Conversely, Belinuridae have more elongate cephalothoracic appendages relative to body length. Differences in appendage sizes are likely related to the freshwater and possible subaerial life modes of belinurids, contrasting with the primarily marine habits of other families. The morphological similarity of T. syriacus to extant members of the genus indicates that the conserved nature of the generic lineage can be extended to ecological adaptations, notably burrowing, swimming, possible diet, and sexual dimorphism.

MORPHOLOGICAL ASSESSMENT OF HORSESHOE CRABS (TACHYPLEUS TRIDENTATUS) BY USING MAGNETIC RESONANCE IMAGING.

The population of Tachypleus tridentatus, a horseshoe crab, has decreased sharply in Japan. With the exception of a few reports regarding the morphological features of T. tridentatus, a three-dimensional analysis of its inner anatomical structures has never been documented. Here, the anatomical structures in two T. tridentatus specimens were examined using magnetic resonance imaging (MRI). In the larger of the two specimens, the three-dimensional T2-weighted image clearly revealed muscles, digestive and urinary systems, ovary, and brain; however, the cardiovascular system was poorly visualized. In the smaller specimen, cardiovascular, digestive, and urinary systems were identifiable. In the three-dimensional T1-weighted images, tissue contrast was low, making it challenging to distinguish the individual organs. These results suggest that MRI is a powerful tool for visualizing the inner anatomical structures of horseshoe crabs.

Computational biomechanical analyses demonstrate similar shell-crushing abilities in modern and ancient arthropods.

The biology of the American horseshoe crab, Limulus polyphemus, is well documented-including its dietary habits, particularly the ability to crush shell with gnathobasic walking appendages-but virtually nothing is known about the feeding biomechanics of this iconic arthropod. Limulus polyphemus is also considered the archetypal functional analogue of various extinct groups with serial gnathobasic appendages, including eurypterids, trilobites and other early arthropods, especially Sidneyia inexpectans from the mid-Cambrian (508 Myr) Burgess Shale of Canada. Exceptionally preserved specimens of S. inexpectans show evidence suggestive of durophagous (shell-crushing) tendencies-including thick gnathobasic spine cuticle and shelly gut contents-but the masticatory capabilities of this fossil species have yet to be compared with modern durophagous arthropods. Here, we use advanced computational techniques, specifically a unique application of 3D finite-element analysis (FEA), to model the feeding mechanics of L. polyphemus and S. inexpectans: the first such analyses of a modern horseshoe crab and a fossil arthropod. Results show that mechanical performance of the feeding appendages in both arthropods is remarkably similar, suggesting that S. inexpectans had similar shell-crushing capabilities to L. polyphemus This biomechanical solution to processing shelly food therefore has a history extending over 500 Myr, arising soon after the first shell-bearing animals. Arrival of durophagous predators during the early phase of animal evolution undoubtedly fuelled the Cambrian 'arms race' that involved a rapid increase in diversity, disparity and abundance of biomineralized prey species.

A 3D anatomical atlas of appendage musculature in the chelicerate arthropod Limulus polyphemus.

Limulus polyphemus, an archetypal chelicerate taxon, has interested both biological and paleontological researchers due to its unique suite of anatomical features and as a useful modern analogue for fossil arthropod groups. To assist the study and documentation of this iconic taxon, we present a 3D atlas on the appendage musculature, with specific focus on the muscles of the cephalothoracic appendages. As L. polyphemus appendage musculature has been the focus of extensive study, depicting the muscles in 3D will facilitate a more complete understanding thereof for future researchers. A large museum specimen was CT scanned to illustrate the major exoskeletal features of L. polyphemus. Micro-CT scans of iodine-stained appendages from fresh, non-museum specimens were digitally dissected to interactively depict appendage sections and muscles. This study has revealed the presence of two new muscles: one within the pushing leg, located dorsally relative to all other patella muscles, and the other within the male pedipalp, located in the modified tibiotarsus. This atlas increases accessibility to important internal and external morphological features of L. polyphemus and reduces the need for destructive fresh tissue dissection of specimens. Scanning, digitally dissecting, and documenting taxa in 3D is a pivotal step towards creating permanent digital records of life on Earth.

Exceptional appendage and soft-tissue preservation in a Middle Triassic horseshoe crab from SW China.

Horseshoe crabs are classic "living fossils", supposedly slowly evolving, conservative taxa, with a long fossil record back to the Ordovician. The evolution of their exoskeleton is well documented by fossils, but appendage and soft-tissue preservation is extremely rare. Here we analyse details of appendage and soft-tissue preservation in Yunnanolimulus luopingensis, a Middle Triassic (ca. 244 million years old) horseshoe crab from Yunnan Province, SW China. The remarkable preservation of anatomical details including the chelicerae, five pairs of walking appendages, opisthosomal appendages with book gills, muscles, and fine setae permits comparison with extant horseshoe crabs. The close anatomical similarity between the Middle Triassic horseshoe crabs and their recent analogues documents anatomical conservatism for over 240 million years, suggesting persistence of lifestyle. The occurrence of Carcinoscorpius-type claspers on the first and second walking legs in male individuals of Y. luopingensis indicates that simple chelate claspers in males are plesiomorphic for horseshoe crabs, and the bulbous claspers in Tachypleus and Limulus are derived.

Augmented laminography, a correlative 3D imaging method for revealing the inner structure of compressed fossils.

Non-destructive imaging techniques can be extremely useful tools for the investigation and the assessment of palaeontological objects, as mechanical preparation of rare and valuable fossils is precluded in most cases. However, palaeontologists are often faced with the problem of choosing a method among a wide range of available techniques. In this case study, we employ x-ray computed tomography (CT) and computed laminography (CL) to study the first fossil xiphosuran from the Muschelkalk (Middle Triassic) of the Netherlands. The fossil is embedded in micritic limestone, with the taxonomically important dorsal shield invisible, and only the outline of its ventral part traceable. We demonstrate the complementarity of CT and CL which offers an excellent option to visualize characteristic diagnostic features. We introduce augmented laminography to correlate complementary information of the two methods in Fourier space, allowing to combine their advantages and finally providing increased anatomical information about the fossil. This method of augmented laminography enabled us to identify the xiphosuran as a representative of the genus Limulitella.

Simple Eyes, Extraocular Photoreceptors and Opsins in the American Horseshoe Crab.

The eyes and photoreceptors of the American horseshoe crab Limulus polyphemus have been studied since the 1930s, and this work has been critical for understanding basic mechanisms of vision. One of the attractions of Limulus as a preparation for studies of vision is that it has three different types of eyes-a pair of later compound, image-forming eyes and two types of simple eyes, a pair of median ocelli, and three pair of larval eyes. Each eye type is tractable for experimentation. Limulus also has extraocular photoreceptors in its segmental ganglia and tail. The current contribution focuses on photoreceptors in Limulus larval eyes and ocelli and its extraocular photoreceptors with the goal of summarizing what is currently known and not known about their physiology and function and the opsins they express. The Limulus genome encodes a surprisingly large number of opsins (18), and studies of their expression pattern have raised new questions about the role of opsin co-expression, the functions of peropsins expressed outside of eyes, and the physiological relevance of opsins with apparently very low expression levels. Studies of opsin expression in Limulus lead one to wonder whether photoreceptors yet to be discovered might be present throughout its central nervous system.

Population structure of the red mangrove crab, Goniopsis cruentata (Decapoda: Grapsidae) under different fishery impacts: Implications for resource management / Estructura de la población del cangrejo rojo de manglar, Goniopsis cruentata (Decapoda: Grapsidae) bajo diferentes impactos de pesca: Implicaciones para la gestión de los recursos

The red mangrove crab, Goniopsis cruentata, influences the recruitment and composition of plant species in the mangrove ecosystem and it is an important fishery resource. Nevertheless, no current management and conservation plans are available for this species for the Brazilian coast. This investigation evaluated the population structure and reproductive biology in populations of G. cruentata under contrasting fishery pressures. The sampling program was carried out in two mangroves, Vaza-Barris and Sergipe River, from January through December 2011. Crabs from both mangroves were randomly collected by a professional fisherman during daytime low tide periods, using a fishing rod baited with pieces of a locally abundant gastropod, Pugilina morio, during 20min/area (catch per unit effort). Monthly measurements of air, sediment surface layer and water temperatures were obtained with a digital thermometer and salinity with an optical refractometer. Both crab populations were compared concerning their abundance, body size, sex ratio, size at onset of sexual maturity and fecundity (FI). Abiotic factors (air, water and mud temperature; and salinity) showed no significant differences between sampling localities. A total of 4 370 crabs were sampled, 2 829 from the Sergipe River and 1 541 from the Vaza-Barris River. The abundance and body size of crabs were compared between mangroves, and statistically significant differences were found. The sex ratio for both populations differed from the expected 1:1 ratio, and a significant deviation in favor of juvenile males was obtained, while adults showed a bias toward females. The estimated size at onset of sexual maturity for both sexes was similar in both populations. However, the populations differed significantly in the number and volume of eggs: a higher FI was obtained in females from the Sergipe River, while a higher egg volume was observed in females from the Vaza-Barris River mangrove. These results indicated a tendency to decrease the body size, the abundance of crabs and the reproductive potential of the species, with higher fishing intensities, reinforcing the need to develop a management plan as well as to establish conservation units for G. cruentata in the Northeastern Brazilian coast. Habitat loss in conjunction with long-term overfishing can have irreversible consequences, which can impact not only the populations of commercially exploited crabs, but the dynamics of virtually the entire mangrove ecosystem.

El cangrejo rojo Goniopsis cruentata influye en el reclutamiento y la composición de especies vegetales del ecosistema manglar y es un importante recurso pesquero. Sin embargo hasta la fecha, no se han desarrollado planes de manejo y conservación para esta especie en Brasil. La presente investigación evaluó la estructura poblacional y la biología reproductiva de las poblaciones de G. cruentata expuestas a diferentes presiones de pesca. El muestreo se realizó en dos manglares, Río Sergipe y Río Vaza-Barris entre enero y diciembre 2011. En ambos manglares, los animales fueron recolectados de día durante la marea baja por un pescador profesional, utilizando una caña de pescar con carnada del gasterópodo Pugilina morio, durante 20min/área (captura por unidad de esfuerzo, CPUE). Mensualmente con un termómetro digital se registró la temperatura del aire, agua y capa superficial del sedimento, mientras que la salinidad fue cuantificada utilizando un refractómetro óptico. Las poblaciones fueron comparadas en su abundancia, tamaño corporal, proporción de sexos, talla de primera madurez sexual y fecundidad. Los factores abióticos(temperatura del aire, agua y sedimento y la salinidad) no mostraron diferencias significativas entre ambos sitios de muestreo. Se recolectaron un total de 4 370 cangrejos; 2 829 en el río Sergipe y 1 541 en Vaza-Barris. Se registraron diferencias significativas entre los manglares con respecto a la abundancia y el tamaño corporal de los animales. La proporción de sexos de ambas poblaciones difirió de la relación 1:1 esperada y se obtuvo una desviación significativa en favor de machos jóvenes, mientras que los adultos mostraron una tendencia hacia las hembras. A su vez, las poblaciones difirieron significativamente en el número de huevos, con un mayor IF para las hembras del manglar del río Sergipe. El volumen de los huevos también mostró diferencias significativas entre ambas poblaciones. Estos resultados indican que la alta intensidad de pesca podría estar afectando dichos parámetros, lo que refuerza la necesidad de desarrollar un plan de manejo y la creación de unidades de conservación para G. cruentata en la costa noreste de Brasil. La pérdida del hábitat junto con la sobrepesca a largo plazo, pueden tener consecuencias irreversibles, que afectan no sólo a las poblaciones de cangrejos de importancia comercial, sino también la dinámica del ecosistema de manglar.

An "ancient" complexity? Evolutionary morphology of the circulatory system in Xiphosura.

Horseshoe crabs (Xiphosura) have been an object of zoological research for almost 200 years. Although some morphological work on the circulatory system has been done, the three-dimensional structure of this complex organ system has never been shown satisfactorily and some crucial questions remain unanswered. Here, the circulatory systems of juveniles of the horseshoe crab taxa Limulus polyphemus and Carcinoscorpius rotundicauda were investigated using a combination of an injection method and micro-computed tomography. Data were processed and 3D-visualized using reconstruction software. Furthermore, the heart was examined using scanning electron microscopy. Additionally, the histology of some structures was investigated via light microscopy and transmission electron microscopy. The results show the high degree of complexity of the arterial and lacunar systems of Xiphosura and provide insights into their three-dimensional structure and relationship to other organ systems such as the central nervous system. We show that the major lacunae, previously described as vessel-like - though indeed highly ramified - can clearly be distinguished from arteries in histological sections because they have no distinct walls. Similarities and differences between the xiphosuran species and arachnids are highlighted and possible phylogenetic implications and evolutionary scenarios discussed.

The horseshoe crab of the genus Limulus: living fossil or stabilomorph?

A new horseshoe crab species, Limulus darwini, is described from the uppermost Jurassic (ca. 148 Ma) near-shore sediments of the Kcynia Formation, central Poland. The only extant species Limulus polyphemus (Linnaeus) inhabits brackish-marine, shallow water environments of the east coast of the United States. Here it is shown that there are no important morphological differences between the Kcynia Formation specimens and extant juvenile representatives of the genus Limulus. The palaeoecological setting inhabited by the new species and the trophic relationships of extant horseshoe crabs are discussed in an attempt to determine the potential range of food items ingested by these Mesozoic xiphosurans. In this paper we propose the adoption of a new term stabilomorphism, this being: an effect of a specific formula of adaptative strategy among organisms whose taxonomic status does not exceed genus-level. A high effectiveness of adaptation significantly reduces the need for differentiated phenotypic variants in response to environmental changes and provides for long-term evolutionary success.