Substrate specificity in the Devonian tabulate coral Pleurodictyum

The tabulate coral Pleurodictyum americanum Roemer has been cited as an example of a host-specific organism occurring exclusively on the shells of gastropods, particularly Palaeozygopieura hamiltoniae (Hall). Examination of over 1600 specimens of P. americanum, from the Middle Devonian Hamilton Group of western New York, reveals additional complexities which require reinterpretation. While substrate selectivity for Palaeozygopieura shells is evident in all 42 subsamples, a variety of other substrates were also utilized by Pleurodictyum including corals, brachiopods, other molluscs and pebbles. Recent scleractinian corals inhabiting soft bottoms show similar substrate preference, selecting for the tubes of live serpulids, or gastropod shells (invariably with a secondary sipunculid host), but also occasionally settling on unoccupied shells or pebbles. Shell surfaces of P. hamiltoniae, preserved as external molds on the Pleurodictyum epitheca, exhibit encrustation by worm tubes and bryozoans as well as borings and mechanical shell damage, suggesting that these were not the shells of live gastropods. However, the invariant aperture-downward orientation and the high degree of selectivity of P. americanum strongly suggest that the shells were occupied by secondary hosts. □ Substrate specificity, commen-salism, tabulate coral, gastropod, sipunculid, Devonian, Hamilton Group, New York.     

Crab shell-crushing predation and gastropod architectural defense

The shell-breaking behavior of the crabs Ozius verreauxii Saussure 1853 and Eriphia squamata, Stimpson 1859 from the Bay of Panama is described. The master claws of both these crabs are well designed for breaking shells. Small shells, relative to the size of a crab predator, are crushed by progressively breaking off larger segments of a shell's apex, while larger shells are peeled by inserting a large dactyl molar into the aperture of a shell and progressively chipping away the lip of the shell.

Heavy gastropod shells are shown to be less vulnerable to crab predators than lighter shells, and narrow shell apertures and axial shell sculpture are demonstrated to be architectural features that deter crab predation. The incidence of architectural features which deter crab predation appears to be higher for smaller gastropod species than for larger gastropods which are too large for most crab predators. Large fish predators prey upon both gastropods and shell-crushing crabs. To avoid fish predators, both these prey groups seek refuge under rocks when covered by the tide. Fish predation thus appears to enforce a close sympatry between smaller gastropods and their crab predators.     

The “<Emphasis Type="Italic">inoceramus</Emphasis> limpet”<Emphasis Type="Italic">Gigantocapulus problematicus</Emphasis> (<Emphasis Type="SmallCaps">Nagao &; Otatume</Emphasis>, 1938) in New Zealand (Late Cretaceous Gastropoda or Monoplacophora,Gigantocapulidae n. fam.)

Limpets up to at least 200 mm long occur widely in New Zealand early Haumurian (Campanian) rocks. Three specimens are in life position, attached to articulated specimens ofIsognomon. The limpet has several unique characters — an apparently anterior marginal apex (all gastropod limpets have a central or posterior apex), an enormously thickened apical area, and a tongue-like projection, with a gape at each side, below the margin at the anterior — demonstrating that it belongs inGigantocapulus Hayami &; Kanie. The low shape, weak sculpture, and marginal apex indicate that it belongs inG. problematicus (Nagao &; Otatume), previously reponed only from Japan and Kamchatka. New Zealand material supports previous interpretations that the apex was anterior, and thatGigantocapulus was an epiparasite or, more probably, a filter feeder living sedentarily on bivalves. The shell is, uniquely, composed of calcite in multiple complex crossed-lamellar layers, very different from the aragonite shell with inner nacreous and outer prismatic layers of both early Palaeozoic and present-day tergomyans. Nevertheless, as the anteriori?) tongue-like projection, thickening and gapes closely resemble those of early Palaeozoic tergomyans, no other limpet-shaped molluscs are known with an anterior apex, and not all modem monoplacophorans are minute (Neopilina reaches at least 40 mm long), a position in Monoplacophora (= Tergomya) remains a possibility. However,G. giganteus (Schmidt) has a subcentral apex and is clearly cyclomyan rather than tergomyan.Gigantocapulus is probably a vanikoroidean gastropod, but could also be a tergomyan, a helcionelloidan, or a member of another, now extinct group of gastropods. Gigantocapulidae n. fam. is proposed.     

Spiders and harvestmen as gastropod predators

1. Reports are reviewed of gastropod feeding (malacophagy) by spiders and harvestmen. Although the standard textbooks on arachnids recognise the importance of gastropods as prey of harvestmen, none apparently refers to malacophagy by spiders. A review of several hundred papers on spider feeding habits revealed that species from several families kill and devour slugs and snails in the laboratory and/or field. 2. Malacophagy has been reported most frequently among mygalomorph spiders, and can make up a substantial proportion of the diets of some species, however gastropods make up an insignificant percentage of the prey of most araneomorph spiders. The spiders that eat gastropods are species with broad diets composed predominantly of arthropod prey. No species of spider appears to feed exclusively on gastropod prey. 3. Harvestmen from several families have broad diets that often include gastropods. Several species of the family Trogulidae and at least one species of the family Ischyropsalididae [Ischyropsalis hellwigi (Panzer)] are specialised gastropod predators. The trogulids are slender animals that attack the snail through the shell aperture (shell intruders). Ischyropsalis hellwigi, on the other hand, can crush snail shells with its powerful chelicerae (shell breakers). 4. The review highlights apparent convergent evolution by harvestmen and Carabidae of two mutually exclusive morphologies found among gastropod predators. It also suggests that there is an urgent need for systematic studies to be conducted to establish the extent and ecological importance of malacophagy in natural and anthropogenically altered habitats.     

Editor's Note

A new species of Apobaetis is described based on nymphs and male imago, and the male imago of A. fiuzai. Specimens were collected in the state of Amazonas, northern region of Brazil. Apobaetis hamadae sp. n. can be easily distinguished from the other congeners by the following combination of characteristics: male imago: (1) abdominal colour pattern; (2) area between forceps not excavated medially. Nymphs: (1) distal margin of labrum with shallow medial emargination and with four to five spatulated setae near midline; (2) maxillary palp 1.4× longer than galea-lacinia; (3) tarsal claws 1.2× times longer than tarsus. The male imago of A. fiuzai can be easily distinguished by the following characteristics: (1) abdominal colour pattern; (2) area between segments I of forceps with a deep V emargination.     

寒武纪早期澄江生物群中受创伤的中间型始莱德利基三叶虫(Eoredlichia intermedia)

三叶虫是寒武纪海洋中的常见动物,其受伤愈合现象的化石记录可以反映生态系统内物种之间竞争关系。中国华南寒武纪早期澄江化石生物群因保存软躯体构造化石而成为研究寒武纪大爆发时期古海洋生态群落的绝佳窗口,然而,澄江生物群中三叶虫受伤愈合现象的化石至今未见报道。中间型始莱德利基虫(Eoredlichia intermedia Lu,1940)是澄江生物群的常见物种,也是我国寒武系第二个三叶虫化石带的标准带化石。在2 000余枚中间型始莱德利基虫标本中,发现两枚成体标本的头甲侧边缘、颊刺和胸甲肋节保存了不同程度的残缺。据此推测,Eoredlichia intermedia在生命活动过程中曾受到碎壳型捕食动物的攻击或其他伤害而形成伤口。伤口部位边缘加厚且光滑,显示了明显的愈合痕迹。这些特征表明始莱德利基虫在受到非致死的伤害后,具备自身修复损伤的能力。这是迄今所知最为古老的,也是在澄江生物群中首次发现的三叶虫受伤并且愈合的标本,反映了寒武纪第二世第三期底栖生态系统中碎壳型捕食者和三叶虫猎物之间的军备竞赛生态关系。     

Evidence of predation in Early Cretaceous unionoid bivalves from freshwater sediments in the Cameros Basin,Spain

Here, we present evidence of possible vertebrate predation on freshwater bivalves from the Lower Cretaceous strata of the Cameros Basin (Spain). The described collection contains the largest number of vertebrate‐inflicted shell injuries in freshwater bivalve shells yet reported in the Mesozoic continental record. Several types of shell damage on fossil shells of Protopleurobema numantina (Bivalvia: Unionoida) are described and their respective modes of formation interpreted in the context of morphological attributes of the shell injuries and the inferred tooth morphology of predators that could have inflicted such injuries. Detailed study of these bite marks shows similarities with the well‐documented injuries in the shells of marine molluscs, namely ammonoids, that have likewise been attributed to reptilian predators. The most parsimonious interpretation suggests crocodiles as the vertebrates interacting with the bivalves in the Cameros Basin. □Barremian–Aptian; bite marks; freshwater bivalves; predation; reptile; Unionoida.     

Spines and epibionts as antipredator defenses in the thorny oyster Spondylus americanus Hermann

Observations and experiments on the cemented pelecypod Spondylus americanus Hermann at Discovery Bay, Jamaica were made to identify the predators of this animal and learn whether their spines, or their epibionts, are important in deterring predation. Spine growth was studied also and found to be very rapid (≈ 1mm/day), and broken spines broken were repaired at this rate.The predators of Spondylus include the gastropod Fasciolaria tulipa Linnaeus (responsible for half of the observed predation), and various crushing predators, including the spiny lobster [Panulirus argus (Latreille)], rays [Aetobatis narinari (Euphrasen) and Dasyatus americanus Hildebrand], the porcupine fish (Diodon hystrix L.), and an unidentified stomatopod. Predators show an increased frequency of attacks in shallow water versus deep, and probably account for the scarcity of Spondylus americanus above −15 m.The concealment effect of epifauna was found to be more important than spines as an antipredator defense. Removal of spines and epibionts significantly increased levels of mortality only when both features were manipulated. There was no statistically significant increase in predation following the removal of epibionts because the shells were rapidly grown over by sponges. Spines alone do not appear to deter predators, but probably serve to attract epibionts and extend them beyond the more vulnerable shell margin. These results do not support the view that spondylid spines evolved as structural deterrents to predators. I argue that they most likely evolved to attract other organisms to settle on the shell and conceal it from predators.     

Drilling predation on scaphopods and other molluscs from the Upper Cretaceous of Manitoba, Canada

Abundant information on drilling predation upon fossil bivalves, gastropods, and brachiopods has been accumulated during the past several decades, but very little is known about the predation on marine, infaunal carnivorous scaphopods. A survey of over 440 specimens of the scaphopod Dentalium gracile collected from the Upper Cretaceous Millwood Member (Campanian) of the Pierre Shale at a site near Russell, Manitoba displays a drilling percentage of approximately 35%. This figure is higher than previously reported for the scaphopods of similar age elsewhere, but is comparable to or lower than that of the sub-Holocene (34–61%). The difference in drilling percentage among different collections may be taxon-related or affected by the composition and structure of the fossil community. Morphologically, the drill-holes, produced by predatory drilling, are beveled with a countersunk profile of clean sharp edges. The drill-hole inner margin is ovate whereas the outer margin is nearly circular. Among the Manitoba specimens, drill-holes seem to be more common on the lateral side. Presumably due to the lack of effective defense strategy, the prey effectiveness is low (∼3.2%). There is no correlation between drill-hole dimension and scaphopod prey size, indicating that predator size does not correlate with the prey size. Nearly 480 specimens of cephalopods, pelecypods, and gastropods were also collected from the same community. There were only a total of 16 drill-holes on this additional material. Over 400 specimens of the only naticid Euspira obliquata were recovered from the same site and are believed to be the predators of the scaphopods and other molluscs. The low percentage (∼2.5%) of drilling on the abundant gastropod E. obliquata may indicate mistaken or opportunistic attacks by the predator, or that the cannibalism is not common in this community.     

Shell repair and mode of life of Praenatica gregaria (Gastropoda) from the Devonian of Bohemia (Czech Republic)

Repaired shell injuries occuring in less than four per cent of examined specimens of the platyceratid gastropod Praenatica gregaria (Barrande, in Perner 1903) from the Lower Devonian (Pragian) of Bohemia probably result from non–lethal predatory attacks. The low frequency may reflect the resistance of the large, smooth, rapidly expanding shell to attack, but a high rate of fatal attacks could produce the same pattern. The habit of some platyceratid gastropods living on echinoderm calices may provide a refuge from benthic predators, but conclusive evidence for this mode of life in P. gregaria or the general assumption of platyceratid coprophagy is lacking.     

Antipredatory traits of the ammonoid shell — Indications from Jurassic ammonoids with sublethal injuries

More than 1500 Jurassic ammonoids with sublethal injuries from the H.Keupp collection (Berlin) were examined. They came from the Toarcian of Southern France, Southern Germany and England, from the Callovian of Villers-sur-Mer (Normandy, France) and from the Oxfordian of Sacaraha (Madagascar). Most sublethal injuries observed can not be referred to specific predators but display the intrinsic factors like tolerance against irritations and disturbances caused by predatory attacks. The types of shell breakages are strongly dependent on shell shape and specific sculpture. Some of the observed types occur exclusively in specific morphotypes. Average range of observed injuries is highest in weakly ornamented shells and lowest in highly ornamented shells. Maximum observed aboral range of shell breakages occurred in longidomes, the lowest in brevidomes and mesodomes. Shell form, sculpture and the length of the living chamber influence susceptibility to lethal injuries, confirmed by the frequency of multiple injuries, which is apparently a good proxy of the vulnerability towards shell crushing predators. The antipredatory traits were in an adaptational conflict with traits demanded for high manoeuvrability and streamlining.      

A compsemydid turtle from the Upper Cretaceous of Var,southern France

Calissounemys matheroni gen. et sp. nov. (Testudines) is described on the basis of a skull and shell elements from the Upper Cretaceous of Var, southern France. This new taxon is assigned to the family Compsemydidae and characterized by a thick-boned, robust skull, a shallow temporal emargination, a crista supraoccipitalis not extending beyond the posterior edge of the skull roof, large nasals meeting along the midline for their full length; frontals retracted from the orbital margin, absence of a cheek emargination, a large jugal forming a substantial part of the orbital margin, absence of a secondary palate and an uneven upper triturating surface; and the shell with vertebral 1 clearly wider than vertebrals 2-3, with the lateral margins strongly divergent towards the anterior border and wider than long vertebrals 2-3. This find increases the diversity of the Late Cretaceous turtle fauna from southern France, and fills a stratigraphical gap in the fossil record of Compsemydidae between the Early Cretaceous and the Paleocene in Europe.     

The cranial and cervical osteology of the european oystercatcher Haematopus ostralegus L.

The cranial and cervical osteology of the European oystercatcher, Haematopus ostralegus L. is completely described from both whole skeletons and spirit specimens. Contrary to previous reports, the postorbital ligament shows variability in bifurcation and attachment, with the posterior branch, where present, attaching either to the suprameatic process or the zygomatic process. In addition, the quadrate ligament, which had been described as originating from the suprameatic process, appears instead to attach to the zygomatic process in some specimens. The caudal mandibular fenestra, earlier considered absent in oystercatchers, is present in all specimens examined. The cranium is additionally distinguished by a number of features which may be unique to oystercatchers. A fourth, previously unrecorded, division of the maxillopalatine strut is present, while the palatine process of the premaxilla is reduced to a thickening along the edge of the premaxillary process of the palatine. The distinctiveness of the oystercatcher cranium is particularly evident in the area of the quadratomandibular articulation, which possesses features potentially important for stabilizing the joint. A deep retroarticular notch is present, guiding a large medial jugomandibular ligament along the posterior margin of the articulation. A lateral mandibular tuberosity, which is received dorsally by an emargination of the jugal arch and quadrate, may act as an osteological brace, preventing posterior shifting of the closed mandible. The 15 cervical vertebrae are divided on the basis of structural criteria into three sections.     

Symbiosis between gastropods and bryozoans in the late Ordovician of Cumbria, England

Five examples of symbiosis between gastropods and trepostome bryozoans are described from the Ashgill (late Ordovician) of Cumbria (England). The gastropods are invariably found associated with the bryozoans. whereas the bryozoans may be free-living. Encrustation is considered to have taken place, in most cases, on mature, living gastropod shells, resulting in the trepostome utilizing the shell as a surrogate basal disc. Three specimens show encrustation on an empty gastropod shell, causing the trepostome to develop a basal disc similar to non-encrusting forms. The bryozoan colony probably benefited from the symbiotic relationship by increased water flow over the colony, caused by gastropod locomotion, whereas the gastropod was afforded protection from predators. The new species Spiroecus nidhoeggi. Diplotrypa hvergelmi and Monotrypa fontinalis are described.     

Winning the biodiversity arms race among freshwater gastropods: competition and coexistence through shell variability and predator avoidance

Explanations for the coexistence of many closely related species in inland waters continue to be generated more than 50 years after Hutchinson’s question: why are there so many kinds of animals? This review focuses on the hypothesis that high species diversity of freshwater gastropods results, in part, from predators maintaining biodiversity across a range of deep- and shallow-water habitats. Invertebrate predators, such as aquatic insects, and leeches consume soft tissue of pulmonate snails by penetrating shells of various shapes and sizes. Crayfish and large prawns chip around the shell aperture to enter thick shells and crush small shells with their mandibles. Crabs use their strong chelae to crush thin and thick shells. Fishes with pharyngeal teeth are major shell-breaking predators that combine with other vertebrate predators such as turtles and wading birds to increase the diversity of gastropod communities by regulating the abundance of dominant species. Although the generalized diets of most freshwater predators preclude tight co-evolutionary patterns of responses, there are combinations of predators that modify gastropod behavior and shell morphology in aquatic assemblages of different ages and depths. This combination of invertebrate and vertebrate predatory impacts led to competitive advantages among individual gastropods with different adaptations: (1) less vulnerable shell morphologies and sizes; (2) predator-avoidance behaviors; or (3) rapid and widespread dispersal with variable life histories. Some individuals develop thicker and/or narrow-opening shells or shells with spines and ridges. Other thin-shelled species crawl out of the water or burrow to lower their risk to shell-breaking or shell-entering predators. Some alter their age at first reproduction and grow rapidly into a size refuge. Fluctuations in water levels and introductions of non-native species can change competitive dominance relationships among gastropods and result in major losses of native species. Many different gastropod predators control species that are human disease vectors. Most snails and their predators provide other ecosystem services such as nutrient cycling and transfer of energy to higher trophic levels. Their persistence and diversity of native species require adaptive management and coordinated study.     

Operculum regeneration following failed predation in the Silurian gastropod Oriostoma

Regeneration of the calcareous rigiclaudent operculum following severe, non‐lethal, predatory attacks is described in two specimens of the characteristic Silurian gastropod Oriostoma from the English Midlands. This is the first record of the regeneration of opercula in Palaeozoic gastropods and of opercula conjoined with shells that have been repaired after significant, failed, durophagous attacks. In one specimen, a series of repaired injuries culminates in a repaired aperture closed by the operculum. In the second specimen, new shell growth has failed to recover the full extent of the original broken whorl, and the new aperture and operculum, though well developed, are smaller than the originals. Both opercula are unusually thin centrally when compared to other, strongly domed, Oriostoma opercula.     

Symbiosis of sea anemones and hermit crabs: different resource utilization patterns in the Aegean Sea

The small-scale distribution and resource utilization patterns of hermit crabs living in symbiosis with sea anemones were investigated in the Aegean Sea. Four hermit crab species, occupying shells of nine gastropod species, were found in symbiosis with the sea anemone Calliactis parasitica. Shell resource utilization patterns varied among hermit crabs, with Dardanus species utilizing a wide variety of shells. The size structure of hermit crab populations also affected shell resource utilization, with small-sized individuals inhabiting a larger variety of shells. Sea anemone utilization patterns varied both among hermit crab species and among residence shells, with larger crabs and shells hosting an increased abundance and biomass of C. parasitica. The examined biometric relationships suggested that small-sized crabs carry, proportionally to their weight, heavier shells and increased anemone biomass than larger ones. Exceptions to the above patterns are related either to local resource availability or to other environmental factors.     

Sublethal injuries in Middle Jurassic ammonite shells from Poland

Sublethal injuries, the effects of which are seen as regeneration patterns, are described from Late Bajocian and Bathonian (Middle Jurassic) ammonites from Poland (Polish Jura area) for the first time. The total number of ammonite shells bearing signs of sublethal injuries is small (only 11 specimens, which constitute ∼1.2% of all ammonites investigated), and this value is even smaller (∼0.3 to 0.8%) when analysing a large sample of a particular ammonite species. Specimens under consideration represent ten species, belonging to six genera and five families. All the healed injuries are represented by only one type, referred to as the ‘forma verticata’ of Hölder. This type of regeneration, very common in ammonite shells in general, is an effect of a puncture injury of the shell-secreting mantle-epithelia at the apertural margin. Although many different extrinsic (mechanical) factors may be responsible for such healed injuries, here it is most plausible they are an effect of either competitive or predatory activities. Other causes, like collision of the ammonite shells with the substrate in a high-energy environment, are excluded because the sea-bottom was soft and situated below the storm wave-base. From many potential predators inhabiting the Polish Basin during the Bajocian and Bathonian, the most likely to have caused these injuries are other ammonites, belemnites and nautiloids. Crabs, which are cited in the literature as a probable perpetrator of the ‘forma verticata’ injuries, appear unlikely here, as the ammonites under discussion were not purely benthic.     

Apical sensory organ in larvae of the patellogastropod Tectura scutum

The apical sensory organ in veliger larvae of a patellogastropod, a basal clade of gastropod molluscs, was studied using ultrastructural and immunohistochemical techniques. Immediately before veligers of Tectura scutum undergo ontogenetic torsion, the apical sensory organ consists of three large cells that generate a very long apical ciliary tuft, two cells that generate a bilateral pair of shorter ciliary tufts, and a neural ganglion (apical ganglion). Putative sensory neurons forming the ganglion give rise to dendrites that extend to the apical surface of the larva and to basal neurites that contribute to a neuropil. The ganglion includes only one ampullary neuron, a distinctive neuronal type found in the apical ganglion of other gastropod veligers. Serotonin immunoreactivity is expressed by a medial and two lateral neurons, all having an apical dendrite, and also by neurites within the neuropil and by peripheral neurites that run beneath the ciliated prototrochal cells that power larval swimming. The three cells generating the long apical ciliary tuft are lost soon after ontogenetic torsion, and the medial serotonergic cell stops expressing serotonin antigenicity in late-stage veligers. The lateral ciliary tuft cells of T. scutum may be homologs of lateral ciliary tuft cells in planktotrophic opisthobranch veligers. A tripartite arrangement of sensory dendrites, as described previously for veligers of other gastropod clades, can be recognized in T. scutum after loss of the apical ciliary tuft cells.     

Validity of the leiognathid fish,Gazza dentex (Valenciennes in Cuvier and Valencinnes, 1835), with designation of a lectotype, and redescription ofG. minuta (Bloch, 1795)

Gazza dentex (Valenciennes in Cuvier and Valenciennes, 1835), having been synonymized withG. minuta (Bloch, 1795), is redescribed as a valid species.Gazza dentex differs fromG. minuta in having deeper body (43.6–51.4% of standard length [SL] vs. 28.3–46.5% of SL), a broad anterodorsal extension of subocular silvery region, in contact with orbit proximally and distally (vs. a long narrow anterodorsal extension, proximal contact only with orbit), scaled area of anterior dorsolateral suface of body not beyond a vertical through posterior tip of sensory canal on temporal (vs. beyond), distance from posterior margin of temporal to anterior tip of dorsolateral scaled area equal to length of 3–5 anterior pored lateral line scales (vs. length of 1–2.5 anterior pored lateral line scales), some dark narrow wavy bands dorsolaterally on body (vs. some dark broad wavy bands above lateral line and a row of dark spots along lateral line), first interneural inserted deeply between first and second neural spines (vs. inserted shallowly), anterior expansion of first interneural narrow, its margin concave (vs. anterior expansion broad, its margin broadly convex), antrorse extension of first interhemal short, deep, acutely pointed (vs. long, moderately deep, pointed) first to fourth hypurals forming 2 plates (first+second and third-fourth hypurals) (vs. a single plate). The lectotype and three paralectotypes are designated forG. dentex, andG. minuta is redescribed.