Reef framework diagenesis across wave-flushed oxic-suboxic-anoxic transition zones

 The temporal and spatial variability of inorganic nutrient concentrations in overlying- and interstitial-seawater in Checker Reef, Oahu was examined for response to incident wave magnitude and direction. Well-point samplers were used to profile interstitial nutrient concentrations across oxic-suboxic-anoxic transition zones in the upper meter of the reef framework at four sites aligned across the patch reef. Samples were acquired over February, 1992, during which time dominant E-NE trade winds directed waves across the reef from the fore-reef to back-reef. However, W-SW “Kona” winds periodically interrupted this pattern and directed waves in the reverse direction. The interstitial microbial habitats of fore- and back-reef framework were distinct from those within the mid-reef framework. Maximum concentrations of PO₄, Si, and NH₄ in interstitial waters occurred at framework depths of 1–2 m, with the highest concentrations occuring within the mid-reef framework. Maximum concentrations of NO₃ and NO₂, which were used to delineate the core of the suboxic zone, occurred at framework depths of 5–10 cm at all stations and attained 2–4 fold higher peak concentrations within the mid-reef and back-reef than within the fore-reef. Variability in interstitial nutrient concentrations was greatest within the back-reef and is consistent with reversals of wave-direction, with the resultant increases in mixing between interstitial and overlying seawater due to flushing caused by the S-SW Kona wind events. The ratio of molar concentrations of total inorganic nitrogen to phosphate (TIN : PO₄) for the fore-reef was 5 : 1; while ratios for the mid- and back-reef were 13–15 : 1, reflecting that the dominant source of particulate organic matter to the fore-reef framework is plankton, while that of the mid- and back-reef is benthic reef plants. Accepted: 4 May 1999     

Paleoecology of the cenozoic reefal foraminifers and algae — A brief review

The paleoecology of reefal foraminifers and algae assumes a considerable importance in determining and delineating sub-environments of ancient reefs, especially those of non-coral origin.A review of the ecologic distribution of the Cenozoic larger foraminifers in different biofacies of the reef-complex environment has revealed the following: (1) a prolific growth of “Alveolina” was possible in the back-reef region near the reef core; (2) Orbitolites and Marginopora preferred sheltered waters on the reef-flat and in the back-reef zones; (3) nummulitids and Discocyclina thrived in both fore- and back-reef shoal areas, but the species living in the former are much stouter than those living in the latter; (4) Heterostegina is and, in the geologic past, was a form, preferring quieter waters of the back-reef lagoons and reef-flat pools; (5) Pellatispira was a typical fore-reef form.Smaller foraminifers, as a whole, are dominant in back-reef lagoons. An abundance of miliolids indicates a sheltered environment prevailing in the reef-flat pools and back-reef zones, whereas reef flats, in general, are characterized by a paucity of smaller foraminifers. An increase in the number of nodosariids and globigerinids points to a fore-reef environment, the depth of which is indicated by the relative abundance of the latter group. Encrusting foraminifers are characteristic of the reef core and are important constituents of for-algal (foraminiferal + algal) reef complexes.Of the algae, the calcarous chlorophyte Halimeda is relatively more abundant in the sheltered parts of a reef-complex, especially the lagoons, where water is moderately agitated and clear; its sudden abundance in the geologic record indicates the advent of a reefal environment. An abundance of the calcareous chlorophyte Dasycladaceae indicates the shallow back-reef areas adjacent to the reef core. Articulated coralline algae are associated with reef-complexes but are varied in their adaptability and, hence, are widely distributed in different parts of the complex. Abundant crustose coralline algae almost certainly indicate a reef-core sub-environment; their skeletons are among the chief constructional units of the core. They increase in abundance towards the outer edge of the reef core and decrease away from it.     

Plant defense associations in the marine environment

In contrast to terrestrial systems, few positive plant-plant associations have been recorded in tropical reef environments. This study, conducted at Carrie Bow Cay, Belize during 28 March–10 April 1984, provides the first documentation of herbivore escapes for natural combinations of palatable and unpalatable marine plants. For example, there was a highly significant association of several macrophyte taxa (Laurencia poitei, Dictyota spp., Amphiroa fragilissima, Cladophoropsis macromeres, Galaxaura cylindrica, rhodophycean turf) within a 2.0-cm radius of the herbivore-resistant brown alga Stypopodium zonale. Almost twice as many taxa occurred within 10 cm of S. zonale as within 10 cm of an equal number of random Stypopodium-free points, and there were no algal species negatively associated with S. zonale. The association of A. tribulus, L. poitei, Digenia simplex, rhodophycean turf, and Jania adherens with S. zonale provided them a fourfold greater survivorship per 48 h in the presence of grazing activity by fishes (mainly Acanthuridae and Scaridae). Reduced herbivory by fishes on macroalgae associated with S. zonale was not solely a consequence of its structural aspect. Losses of the palatable alga Acanthophora spicifera were significantly greater for thalli spatially remote (30 and 60 cm) from either a real or simulated Stypopodium; however, losses of A. spicifera adjacent to actual Stypopodium plants were significantly less than the losses next to models. The inter-relationships studied here, where an abundant and well-defended plant provides a significant refuge habitat for at least five relatively edible macroalgae, clearly facilitates the survival of certain taxa in the reef system and concomitantly enhances the within habitat diversity. Our findings also suggest an interaction counter to the process of competitive exclusion, since the single predominant plant has a positive rather than negative net effect on the abundances of other species that utilize the same general resources (e.g., light, space, nutrients).     

Macroborers within coral framework at Discovery Bay, north Jamaica: species distribution and abundance, and effects on coral preservation

 Macroboring organisms are recognised as key agents of reef framework modification and destruction, and while recent studies in the Pacific have improved understanding of spatial variations in macroboring community structure, and rates of macroboring within individual reefs, comparable studies from the Caribbean are largely lacking. This study assesses the distribution of macroboring species and the degree of framework infestation across the reefs at Discovery Bay, north Jamaica. Although individual species of borers exhibit variable distributions across the reef, relative abundances of the main groups of macroborers (sponges, bivalves, worms) illustrate clear distributional trends. Sponges are dominant at fore-reef sites, while sipunculan and polychaete worms are only of importance at back-reef/lagoon and shallow fore-reef sites. Bivalves are locally important within back-reef and lagoon patch reef framework. Average percentages of internal bioerosion (macroboring) vary between sites, but are highest at back-reef and deep fore-reef sites. No systematic pattern of variation occurs within back-reef/lagoon samples, but a significant trend of increased macroboring is recognised with increased water depth on the fore-reef. In addition, significant differences in terms of the susceptibility of individual coral species are recognised. These factors are likely to result in biasing of the fossil record, with variable styles of preservation evident both between sites (i.e. with depth/environment) and within sites (i.e. between coral species). Accepted: 1 June 1998     

Holocene coral reef rubble and its binding agents

A literature review regarding reef rubble (defined as mechanically or chemically abraded parts of framebuilders or reef rock larger than sand fraction) and its binding agents is presented. Rubble is produced by natural and man-made events such as storms, wave agitation, earthquakes, bioerosion, ship groundings, and dynamite fisheries. The regeneration of reefs after rubble-forming processes requires rigid rubble binding, which is always preceded by preliminary stabilization. Preliminary stabilization can be achieved by a decline in hydrodynamic energy, interlocking of components, seagrass, and overgrowth by sponges or algae. Rigid binding is primarily achieved by diagenetic cementation. The literature indicates that binding by coralline algae or other organisms (corals, worms, bryozoans) is only of subordinate importance. Highest rates of rigid rubble binding are known from fore-reef areas with low sloping angles above fair-weather wave base; rigid rubble binding is particularly rare in deeper fore-reef environments and not described from the reef crest. Rigid binding by diagenetic cementation is generally known from inter- and supratidal near-shore ramparts as well as back-reef, reef-flat, and shallow fore-reef rubble accumulations, while coralline algae rigidly bind rubble only in very shallow fore-reef environments. Rubble binding does not appear to be easily achieved and fewer reports of bound rubble were found than of loose rubble.     

The Ecological Perspective of Microbial Communities in Two Pairs of Competitive Hawaiian Native and Invasive Macroalgae

Marine macroalgae are known to harbor large populations of microbial symbionts, and yet, microbe symbiosis in invasive macroalgae remains largely unknown. In this study, we applied molecular methods to study microbial communities associated with two invasive algae Acanthophora spicifera and Gracilaria salicornia and the two native algae Gracilaria coronopifolia and Laurencia nidifica at spatial and temporal scales in Hawaiian coral reef ecosystems. Bacterial communities of both the invasive and native macroalgae displayed little spatial and temporal variations, suggesting consistent and stable bacterial associations with these macroalgae. Results of this study identified three types of bacterial populations: nonspecific (present in both algal and water samples); algae-specific (found in all algal species); and species-specific (only found in individual species). The bacterial diversity of invasive algae was lower than that of their native counterparts at phylum and species levels. Notably, the vast majority (71 %) of bacterial communities associated with the invasive algae G. salicornia were representatives of Cyanobacteria, suggesting a potential ecological significance of symbiotic Cyanobacteria.     

Spatial heterogeneity of benthic community assemblages with an emphasis on reef algae at French Frigate Shoals, Northwestern Hawai‘ian Islands

Reefs in tropical atoll systems have historically been described on a geomorphic basis, and segregated into loosely defined fore-reef, back-reef, and lagoonal reef zones. However, recent oceanographic monitoring data have shown that physical factors within a single geomorphic zone can vary significantly, calling into question whether benthic communities within a single zone are biologically similar. To determine the amount of benthic variability that may occur in a geomorphic zone, percent cover of benthic organisms was determined at the species level for 28 sites in three geomorphic zones at French Frigate Shoals, Northwestern Hawai‘ian Islands. Multivariate statistical analyses found most windward fore-reef and back-reef sites to be statistically similar, but considerable variation to exist among sites within calmer lagoonal areas. Surveys revealed macroalgae to dominate over scleractinian coral species at the majority of sites in this healthy, subtropical reef system, although select lagoonal areas were dominated by dense coral communities.     

Sponge distribution across Davies Reef,Great Barrier Reef,relative to location,depth, and water movement

Sponge populations were surveyed at different depths in three zones of Davies Reef, a large platform reef of the central Great Barrier Reef. Depth is the major discriminatory factor as few sponges are found within the first 10 m depth and maximal populations occur between 15 m and 30 m on fore-reef, lagoon and back-reef slopes. Reef location is another major factor, with the lagoon containing a significantly different sponge population to either the fore-reef or the back-reef slopes. Physical factors are considered to be the major influences behind these patterns. Physical turbulence is strongest within the first 10 m and apparently limits sponge growth within these shallow zones. Insufficient photosynthetic radiation limits the growth of the sponge population below 30 m depth as many of the species are phototrophic with a dependence on cyanobacterial symbionts for nutrition. Sponge populations on the outer (fore- and back-) reef slopes are comparable with each other but different from those on lagoon slopes where currents are reduced and fine sediment loads are higher. The largest populations occur on the back-reef slope where currents are stronger and there are possibly higher concentrations of organic nutrients originating from the more productive shallow parts of the reef. While there are correlations between sponge populations and environmental parameters, data are insufficient to enable more definitive conclusions to be drawn. Most sponge species are distributed widely over the reef, however, some are restricted to a few habitats and, hence, may be used to characterize those habitats.     

The geological effects of hurricanes on coral reefs and the interpretation of storm deposits

Hurricanes occur in belts 7° to 25° north and south of the equator. Reefs growing in these belts suffer periodic damage from hurricane-generated waves and storm surge. Corals down to 20m depth may be broken and removed, branching colonies being much more susceptible to breakage than upright massive forms. Sand cays may be washed away and former storm ridges may migrate to leeward across reef flats to link with islands. Reef crest and reef front coral debris accumulate as talus at the foot of the fore-reef slope, on submarine terraces and grooves, on the intertidal reef flat as storm ridges of shingle or boulders and isolated blocks of reef framework, as accreting beach ridges of leeward migrating shingle, as lobes and wedges of debris in back-reef lagoons, as drapes of carbonate sand and mud in deep off-reef locations in the fore-reef and lagoonal areas. In addition to the coarse debris deposited, other features may aid the recognition of former hurricane events, including the assemblage of reef biota, its species composition and the structure of the skeletons; graded internal sediments in framework cavities; characteristic sequences of encrusting organisms; characteristic shapes of reef flat microatoll corals; and submarine cement crusts over truncated reef surfaces. The abundance of reef flat storm deposits whose ages cluster around 3000–4000 y BP in certain parts of the world most likely relate to a slight fall in relative sea level rather than an increase in storminess during that period. A higher frequency of storms need not result in more reef flat storm deposits. The violence of the storm relative to normal fair-weather conditions influences the extent of damage; the length of time since the previous major storm influences the amount of coral debris created; the length of time after the hurricane, and before a subsequent storm influences the degree of stabilization of reef-top storm deposits and hence their chances of preservation.     

Coexistence of congeneric spiny lobsters on coral reefs: differential use of shelter resources and vulnerability to predators

The notion, previously generated from laboratory experiments, that the local coexistence of spiny lobsters Panulirus guttatus and Panulirus argus on Caribbean reefs may be promoted by a differential use of shelter resources and/or vulnerability to predators was explored in a coral reef in Mexico. Multiple regressions with data collected on 11 occasions on fixed back-reef and fore-reef sites suggested that the densities of the congener, other crevice-dwellers, and predators did not significantly affect the density of P. argus in either reef zone, or of P. guttatus on the back-reef, where coexistence of both lobster species was greatest. In contrast, there was a significant negative relationship between predators and the density of P. guttatus on the fore-reef, where this species was dominant. Congeneric cohabitation in dens was less than expected by chance, but this pattern may reflect a differential use of shelter resources rather than interspecific competition. P. guttatus was more prevalent in dens over the middle and upper third of the reef profile, and P. argus over the lower and middle third of the reef profile. Whether individuals cohabited with conspecifics, congeners, or resided solitarily, P. guttatus was more prevalent at the walls and/or ceiling and P. argus on the floor of dens. This differential use of shelter resources may be related to the differential vulnerability to predators, which may have promoted local coexistence of these congeners in reef habitats.     

Comparative study of the effects of salinity and UV radiation on metabolism and morphology of the red macroalga <Emphasis Type="Italic">Acanthophora spicifera</Emphasis> (Rhodophyta,Ceramiales)

Increase of harmful radiation to the Earth’s surface due to ozone depletion results in higher exposure to harmful ultraviolet- B radiation (UV), while fluctuations in seawater salinity may alter water density, ionic concentration, nutrient uptake, and osmotic pressure. This study evaluated the effects of salinity and UV on metabolism and morphology of Acanthophora spicifera (M.Vahl) Børgesen. Water with 30 and 37 psu [g(salt) kg^–1(sea water)] was used for experiments during 7 d of exposure to UV (3 h per day). We demonstrated that UV treatment predisposed, irrespective of salinity, A. spicifera to a decrease in its growth rate and cell viability, as well as affected its morphological parameters. After exposure to PAR + UVA + UVB (PAB), samples showed structural changes and damage, such as increasing cell wall thickness and chloroplast disruption. Our results indicate that UV led to dramatic metabolic changes and cellular imbalances, but more remarkable changes were seen in samples exposed to high salinity.     

Holocene growth history of an eastern Pacific fringing reef,Punta Islotes,Costa Rica

Rock and sediment cores reveal that a well-developed fringing reef in Golfo Dulce, Pacific Costa Rica, up to 9 m thick was established on Cretaceous basalt about 5500 y BP. It is presently being smothered with fine sediments and is almost completely dead. This reef is made up of three main facies that are represented by comparable extant reef zones: reef-flat branching coral, fore-reef slope massive coral, and fore-reef talus sediment facies. Reef growth began with the establishment of small patch reefs dominantly formed by the branching coral Pocillopora damicornis. P. damicornis spread across the basalt bench and massive colonies of Porites lobata grew on the outer slopes, eventually blocking the seaward transport of Pocillopora fragments to the fore-reef talus sediments. The reef flourished until 500 years ago. Lower accumulation rates during the past 500 years may be due to deteriorating environmental conditions rather than slower growth after the reef reached sea level. Present-day reef communities are severely degraded with less than 2% living coral cover. The increased turbidity associated with the final stage of degradation of this reef is probably related to human activity on the adjacent shores, including deforestation, mining, and road construction.     

Patterns in the distribution of the crinoid community at Davies Reef on the central Great Barrier Reef

The crinoid community of Davies Reef, a midshelf reef in the central Great Barrier Reef, was systematically sampled in all major crinoid habitats. A total of 294 individuals of 27 species-level taxa was found in 25 sites across the reef. Of these 27 taxa, 20 were confidently assigned to known species. The 25 sitesx27 taxa matrix was subjected to an array of pattern extraction and diagnostic techniques — numerical classification, ordination and minimum spanning trees — to elucidate the structure of the community. These analyses revealed a consistent structure characterized by a species-rich ensemble around the periphery of the reef which was attenuated towards the inside of the reef. This structure contrasts strongly with the patterns seen in other major reef communities, such as hard and soft corals, fish or sponges. In these communities, different parts of the reef are characterized by distinctive sets of species, a depthbased zonation of the communities is evident, and the fore-reef slope typically supports a different ensemble from the back-reef slope. We conclude that the crinoid community offers a significant opportunity to observe the coral reef ecosystem from a different perspective.     

Decadal changes in turbid-water coral communities at Pandora Reef: loss of resilience or too soon to tell?

Coral communities were monitored at Pandora Reef, nearshore Great Barrier Reef from 1981 to 2005 using photography and videography. In the 1980s, regional elevation of land-based nutrients in coastal waters (ca. 2–6 times pre-European levels of early 1800s) did not prevent overall recovery of coral cover and diversity following a sequence of environmental disturbances in the 1970s. However, prospects for a repeat of such resilience following catastrophic mortality from high-temperature bleaching in 1998 and a cyclone in 2000 are not clear. Different coral communities around the reef varied greatly in relation to impacts and recovery. Fore-reef communities dominated by acroporids (fast growing branching and tabular Acropora and foliose Montipora) recovered strongly in the 1980s following apparently severe impacts by cyclone, flood and heat wave disturbances in the 1970s, attaining 60–90% cover by stabilizing rubble and outgrowing macro-algae in <10 years. In the back-reef, by contrast, poritid-dominated communities (massive and finger Porites and columnar Goniopora and Alveopora) had more stable trajectories and smaller impact from recent disturbances: recovery was well underway in 2005. The contrasting trajectories of different parts of the reef reflect differential survival of more persistent versus more ephemeral taxa, notably poritids and acroporids, respectively, both major contributors to framework and cover on reefs globally. A repeat of earlier resilience appears possible in the shallow fore-reef, but unlikely in the deeper fore-reef, which had few viable fragments or recruits in 2005. The main limits on recovery may be (1) reduced supply of coral larvae due to widespread regional losses of coral brood stock and (2) the reduced intervals between disturbances associated with global climate change. The presence of a high abundance of Acroporidae is a major pre-disposing risk factor for climate change impacts.     

Some aspects of the morphology and distribution of Microsoriurn linguaeforrne (Polypodiaceae)

Microsorium linguaeforme is reported for the first time from India. Its rhizome is slender, much elongated with a highly dissected stele and leaf gaps in a single median dorsal row. The characteristic branching pattern is a modification of the common pattern in Polypodiaceae, resulting from the displacement of leaf-associated branches and the precocious development of the most basal secondary branch of each primary branch. The first two or three leaves of juvenile plants have no associated branches; thereafter, abaxially-originating traces t o solitary branches are progressively displaced so that the branch trace is close to the preceding leaf trace. Stomata of adult leaves are Copolomesoperigenous and of juvenile leaves Eupolomesoperigenous. The spores are monolete and with a smooth exine. Spore germination is of the Vitiariatype and prothallial development of the Kauliniatype. It is concluded that M. linguaeforme is closely allied to Leptochilus and is probably parental to it.     

Growth forms and palaeoenvironmental interpretation of stromatoporoids in a Middle Devonian reef, southern Morocco (west Sahara)

Growth forms of well-preserved stromatoporoids, including genera Actinostroma, Stachyodes, and Stromatopora, are described for the first time from the Devonian Sabkhat Lafayrina reef complex of southern Morocco (west Sahara), one of the best exposed Middle-Devonian stromatoporoid-dominated fossil reefs. Three facies types representing the well illuminated fore-reef, reef-core and transition to back-reef facies display the distribution and growth of stromatoporoids in a high latitude setting at 40–50° south of the palaeoequator. Stromatoporoids are largely in growth position and reflect the well-preserved reef architecture. Although outcrops are low topography, the reef's prominent profile is indicated by presence of spur and groove form and a clearly defined reef margin. Stromatoporoids are mostly laminar and domical forms, with little evidence of ragged margins, and indicate normal turbulence shallow waters, with low sediment deposition.     

Tree Branching: Leonardo da Vinci's Rule versus Biomechanical Models

This study examined Leonardo da Vinci''s rule (i.e., the sum of the cross-sectional area of all tree branches above a branching point at any height is equal to the cross-sectional area of the trunk or the branch immediately below the branching point) using simulations based on two biomechanical models: the uniform stress and elastic similarity models. Model calculations of the daughter/mother ratio (i.e., the ratio of the total cross-sectional area of the daughter branches to the cross-sectional area of the mother branch at the branching point) showed that both biomechanical models agreed with da Vinci''s rule when the branching angles of daughter branches and the weights of lateral daughter branches were small; however, the models deviated from da Vinci''s rule as the weights and/or the branching angles of lateral daughter branches increased. The calculated values of the two models were largely similar but differed in some ways. Field measurements of Fagus crenata and Abies homolepis also fit this trend, wherein models deviated from da Vinci''s rule with increasing relative weights of lateral daughter branches. However, this deviation was small for a branching pattern in nature, where empirical measurements were taken under realistic measurement conditions; thus, da Vinci''s rule did not critically contradict the biomechanical models in the case of real branching patterns, though the model calculations described the contradiction between da Vinci''s rule and the biomechanical models. The field data for Fagus crenata fit the uniform stress model best, indicating that stress uniformity is the key constraint of branch morphology in Fagus crenata rather than elastic similarity or da Vinci''s rule. On the other hand, mechanical constraints are not necessarily significant in the morphology of Abies homolepis branches, depending on the number of daughter branches. Rather, these branches were often in agreement with da Vinci''s rule.     

Sedimentary development of a continuous Middle Devonian to Mississippian section from the fore-reef fringe of the Brilon Reef Complex (Rheinisches Schiefergebirge, Germany)

The Brilon-reef complex is one of the biggest Devonian carbonate buildups (~80 km²) of the Rheinisches Schiefergebirge. The Burgberg section is located in the southeastern fore-reef area of the Brilon Reef Complex and exposes a succession of strata (117 m thick), which extends from the Middle Givetian (middle varcus conodont Zone) to the Viséan (bilineatus conodont Zone). Field and microfacies observations led to the definition of nine microfacies that are integrated into a sedimentary model divided into off-reef, intermediate fore-reef, and proximal fore-reef sedimentary domains (SD). The off-reef domain (SD1) is the most distal setting observed and is characterized by fine-grained sediments, dominated by pelagic biota and the local occurrence of gravity-flow deposits. The intermediate fore-reef (SD2) is characterized by a mixture of biota and sediments coming from both deeper-water and shallow-water sources and is influenced by storm and gravity-flow currents. In this domain, Renalcis mound-like structures developed locally. Finally, the proximal fore-reef (SD3) corresponds to the most proximal setting that is strongly influenced by gravity-flow currents derived from the Brilon Reef Complex. The temporal evolution of microfacies in the fore-reef setting of the Burgberg section show five main paleoenvironmental trends influenced by the onset, general development, and demise/drowning of the Brilon Reef Complex. Fore-reef to off-reef lithologies and their temporal changes are from the base to the top of the section: (U1)—fine-grained sediments with large reef debris, corresponding to the initial development of the reef building upon submarine volcaniclastic deposits during the Middle Givetian (middle varcus Zone) and first export of reef debris in the fore-reef setting; (U2)—high increase of reef-derived material in the fore-reef area, corresponding to a significant progradation of the reef from the Middle Givetian to the Early Frasnian (maximum extension of the Brilon Reef Complex to the south, disparilis to the falsiovalis conodont biozones); (U3)—progressive decrease of shallow-water derived material and increase of fine-grained sediments and deep-water biota into the fore-reef setting, corresponding to the stepwise withdrawal of the reef influence; from the Middle to the Late Frasnian (jamieae conodont Zone); (U4)—development of a submarine rise characterized by nodular and cephalopod-bearing limestones extending from the Late Frasnian to the Late Famennian corresponding to the demise and drowning of the Brilon Reef Complex as a result of the Late Frasnian Kellwasser events (upper rhenana and triangularis conodont biozones); (U5)—significant deepening of the Burgberg area starting in the Late Famennian, directly followed by an aggrading trend marked by pelagic shales overlying the nodular limestone deposits.     

A radicle solution: morphology and biomechanics of the Eucalyptocrinites ‘root’ system

Eucalyptocrinites is one the most common and familiar mid‐Palaeozoic crinoids and is the exemplar of a form with dendritic radicular holdfasts. American museums have hundreds of specimens of Eucalyptocrinites holdfasts from the Silurian Waldron Shale of Indiana and Kentucky, USA. The radix (‘root’) system of Eucalyptocrinites can be described as comprised of links (branches) that meet at nodes. Measured values include the x‐ and y‐coordinates of the nodes, the distances of the nodes from the stem, the angles between branches, branch length and branch width. Rose diagrams show clearly that the root systems are not isotropic but have preferred orientations. Branch angles are highly variable, but cluster around 60 degrees. Branch lengths and distal branch widths are relatively constant, but branch width increases variably towards the column. The branching pattern can be modelled as a self‐similar (‘fractal’) structure. Several specimens labelled as Eucalyptocrinites show a distinct fivefold symmetry without branching and very likely represent a different taxon. The Eucalyptocrinites radix system, along with the probably stiff dististele, most likely functioned as a rigid plate that resisted rotational forces due to currents acting on the crown. Upstream radicles experienced tension, whereas downstream roots were compressed. This force distribution may explain the observed anisotropies in radix morphology. The ‘roots’ of Eucalyptocrinites and other crinoids have long been compared with the root systems of plants. Although there are superficial similarities, there are fundamental differences.     

Dual Roles of an Algal Farming Damselfish as a Cultivator and Opportunistic Browser of an Invasive Seaweed

Herbivory is a fundamental process determining reef resilience, and while algal farming damselfishes can help shape benthic assemblages, an understanding of their contribution to areas outside of defended territories is relatively unexplored. Here, we demonstrate how the farming damselfish Stegastes marginatus plays a dual role in benthic structuring by 1) contributing to persistence of the invasive macroalga Acanthophora spicifera within a Hawaiian marine protected area, where the macroalga occurred exclusively inside Stegastes territories, and 2) behaving as an opportunistic browser of the exotic alga outside their territorial borders. Greater than 50% of the biomass of tethered A. spicifera was consumed within one-hour when placed outside Stegastes territories, compared to <5% lost from tethers within territories or herbivore exclusion cages. In situ remote video revealed that tethered A. spicifera located outside territories was grazed primarily by the surgeonfish Acanthurus nigrofuscus (∼68% of total bites) and, surprisingly, by S. marginatus (∼27% of total bites) that left their territories to feed on this resource on 107 occasions during 400 min of filming. Further, for over half of those occurrences where S. marginatus grazed on the tethered macroalga outside of territories, they fed alongside conspecifics and other species, displaying little of the aggressiveness that characterizes this damselfish. These results show that S. marginatus plays a wider role in determining benthic assemblages than previously recognized, acting both as cultivators of a canopy-forming invasive macroalga within their territories, and as opportunistic browsers in neighboring sites. Consequently, S. marginatus can affect benthic species composition across their territory borders. These results provide a rare example of interspecific facilitation of an exotic alga by an indigenous marine fish. Accounting for fish behaviors more broadly is important to further our understanding of ecological processes that shape reef ecosystems to improve management of MPAs that often support extensive farming damselfish populations.