Substratum relationships of some tropical Pacific intertidal gastropods

Among tropical Indo-West Pacific herbivorous intertidal gastropods, high intertidal neritids, and limpets and certain lower intertidal limpets are restricted to a narrow range of substratum types. Littorinids, planaxids, low intertidal neritids, most lower intertidal limpets, and the trochid Monodonta labio tolerate a wide variety of substrata. This pattern of substratum specialization is related to the overall adaptive strategies of the various groups: those with a large area of soft tissue in contact with the substratum (neritids and limpets) often specialize to a particular substratum, especially at high shore levels, while those in which the area of contact between animal and substratum is small do not. High intertidal neritids and limpets able to live on a wide range of substrata tend to be species of limited geographic distribution. It is concluded that, even within a tropically similar group of animals (raspers in this case) relevant dimensions of niche separation depend upon the structure of the environment as well as upon the structure of the animals themselves.     

Ecological studies on a tropical limpet,Cellana radiata. Structural variations in the shell in relation to distribution

Structural variations in the shell of the tropical limpet Cellana radiata (Born) inhabiting the high and low water zones and the exposed and protected surfaces at Palm Beach, Waltair Coast, India, were investigated. Attempts have also been made to study which environmental factors have a marked influence upon the characters of the shell. The results are discussed and are compared with those on temperate climate limpets.     

Life history of the temperate subtidal gastropod Umbonium costatum

The life history and reproductive biology of the trochid sand snail Umbonium costatum (Kiener) were investigated on a subtidal sandy shore in Hakodate Bay, Japan between 1988 and 1991. Female U. costatum mature at 1 yr of age (shell diameter-11 mm), reproduce twice (in June-July and September-October) in successive years, grow to a maximum size (shell diameter=22 mm) at age 8 yr, increase annual fecundity with age from 2000 (age 1 yr) to 37000 (age 8 yr), and show a maximum monthly gonad somatic index of 8% which is constant among ages. In comparison to a previously studied life history of a tropical Umbonium vestiatium, temperate U. costatum shows more sustained growth and a longer life span after maturation. This could be explained by: (1) the optimal size model concerned with resource investment, in gametes (Sebens 1987); and by (2) bet hedging to compensate large variability in larval success at high latitudes. These two hypotheses are not mutually exclusive, but both are based on season-related extremes of environment at high latitudes where the period suitable for reproduction is short.     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Intraspecific variation of sesquiterpene lactones associated to a latitudinal gradient in <Emphasis Type="Italic">Smallanthus macroscyphus</Emphasis> (Heliantheae: Asteraceae)

According to theory, variation in plant secondary metabolism against herbivores is driven by variation in biotic and abiotic conditions interacting with plants genotype to determine the expression of resistance traits. Particularly, it has been long postulated that plants growing along latitudinal gradients experience changes in biotic and abiotic interactions, specifically leading to a decrease of plant toxicity towards the poles. We tested this hypothesis using the asteraceous species Smallanthus macroscyphus. Smallanthus species are known to contain sesquiterpene lactones (STLs), bitter compounds with a broad spectrum of biological activities, including deterrence to herbivores. S. macroscyphus showed a decrease in chemical diversity of STLs when investigating populations growing from the tropical regions to less tropical ones. Populations from lower latitudes were found to be more chemically diverse with enhydrin, uvedalin and fluctuanin as main components, while populations southward were chemically fairly uniform, with polymatin A as the main and largely dominant STL. The STL chemistry of S. macroscyphus is in agreement with the hypothesis that plants of tropical forests have a greater diversity of secondary metabolites when compared to their temperate counterparts.     

云南地区种子植物区系过渡性地理分布格局的群落尺度分析

探索植物区系过渡性的地理分布格局,对于理解植物区系的起源、迁移和分布具有重要意义。本研究利用620个植物群落样方,探讨了云南地区植物区系过渡性的地理分布格局及其生物地理意义。研究结果表明,随着纬度、海拔的升高,温带区系成分所占比重呈显著递增趋势,但在经度梯度上呈递减趋势;热带区系成分所占的比重在地理梯度上则表现为相反的变化趋势。地形格局较好地反映了区系成分的地理分布格局;热带区系成分所占比重与温带区系成分所占比重的比值从南到北呈递减趋势。区系过渡性的低值主要出现在滇西北地区和包括滇西南、滇东南与滇南在内的云南南部地区,而高值主要出现在云南中部地区和中海拔地区,这可能与区系迁移过程和海拔梯度有关。热带区系成分在云南地区基带植被中占主导地位;热带区系成分占优势地位的区域主要集中在低海拔地区,占云南土地面积的60％左右,而温带区系成分占优势地位的区域主要集中在高海拔的北部地区,占土地总面积的40％左右。本研究从基带植被的区系性质和温、热带植物区系成分占优势地位的土地面积的构成证实了云南地区可能是一个“热带山原”的假说。区系过渡性随着年平均温度的升高,呈先增强后减弱的单峰格局。当年均温在13．1℃左右时,出现区系平衡点。相对水平分布而言,云南地区植被的垂直分布可能更值得注意。     

Latitudinal variation in local interactions and regional enrichment shape patterns of marine community diversity

While communities are shaped by both local interactions and enrichment from the regional species pool, we propose a hypothesis that the balance of these forces shifts with latitude, with regional enrichment dominating at high latitudes and local interactions dominating at low latitudes. To test this hypothesis, we conducted a latitudinal-scale experiment with marine epifaunal communities. In four regions of the North Atlantic Ocean and Caribbean Sea, we used mimics of ecosystem engineers to manipulate biogenic structural complexity. We iteratively evaluated diversity patterns of experimental communities up to one year after deployment. Additional data were also collected from one of our tropical sites 2.5 years after initial deployment. As hypothesized, we found a reciprocal latitudinal gradient in the effects of the structurally complex mimics and regional enrichment. In the tropics, local diversity was always higher in association with the mimics than in exposed areas that were more open to predation. This effect was consistent across two spatial scales and beyond the one-year timescale of the experiment. In temperate communities, no consistent effects of the mimics on diversity were observed. However, the proportion of species from the regional species pool that were present at the local scale increased from the tropics to the temperate zone, consistent with the hypothesis that higher-latitude communities may experience greater influence from the regional species pool than communities at low latitudes. This study represents the first large-scale experimental demonstration that suggests that the relative impact of local interactions and regional enrichment on community diversity may depend on latitude.     

Generalists and specialists along a latitudinal transect: patterns of thermal adaptation in six species of damselflies

Tropical organisms colonizing temperate environments face reduced average temperatures and dramatic thermal fluctuations. Theoretical models postulate that thermal specialization should be favored either when little environmental variation is experienced within generations or when among-generation variation is small relative to within-generation variation. To test these predictions, we studied six temperate species of damselflies differing in latitudinal distribution. We developed a computer model simulating how organisms experience environmental variation (accounting for diapause and voltinism) and performed a laboratory experiment assaying thermal sensitivities of growth rates. The computer model showed opposing latitudinal trends in among- and within-generation thermal variability: within-generation thermal variability decreased toward higher latitudes, whereas relative levels of among-generation thermal variability peaked at midlatitudes (where a shift in voltinism occurred). The growth experiment showed that low-latitude species were more thermally generalized than mid- and high-latitude species, supporting the prediction that generalists are favored under high levels of within-generation variation. Northern species had steeper, near-exponential reaction norms suggestive of thermal specialization. However, they had strikingly high thermal optima and grew very slowly over most of the thermal range they are expected to experience in the field. This observation is at present difficult to explain. These results highlight the importance of considering interactions between life history and environmental variation when deriving expectations of thermal adaptation.     

Temperature relationships of some tropical Pacific intertidal gastropods

Tissue and environmental-temperature measurements were made using thermistors in Hawaii, Guam, Palau and Singapore in the summer of 1970. Limpets have a very limited ability to regulate body temperature, and several species possessed tissue temperatures greater than those of the substratum. Some neritids and the trochid Monodonta labio possessed tissue temperatures as much as 2.2° to 2.8°C below substratum temperature. Littorinids attached by a mucus film to the substratum were found to have relatively lower body temperatures than those attached by the foot. Among neritids there is a tendency for high intertidal species to be better temperature regulators than low intertidal species. A very important factor in the temperature regulation of intertidal snails appears to be the extent of the area of contact between the animal and the substratum, which is greatest in limpets and least (negligible) in littorinids. Evaporative cooling and light coloration of the shell play a role in certain species and groups.     

Latitudinal symbiont zonation in <Emphasis Type="Italic">Stylophora pistillata</Emphasis> from southeast Africa

Studies on latitudinal gradients in Symbiodinium diversity on scleractinian corals are largely restricted to warm-water low latitude locations, and it appears that there is a shift in symbiont distributions with increasing latitude. The Symbiodinium assemblages of high latitude coral communities have largely been undocumented despite occupying an important transitional zone between tropical and temperate regions. Using a combination of the internal transcribed spacer region 2 (ITS2) and denaturing gradient gel electrophoresis (DGGE), we assessed the cladal and subcladal variability of Symbiodinium in the widely distributed species Stylophora pistillata along a latitudinal transect in southeast African waters which extended into high latitude locations. All colonies examined belonged to clade C. Six unique ITS2-DGGE banding profiles (designated C_spa to C_spf) were observed, which showed a latitudinal distribution from north to south, most likely a result of a gradient in water temperature and irradiance driven by riverine input in the southern regions. Sequence analysis revealed that all sequences except one did not match previously identified clade C sub-types, probably due to the lack of regional information in the Western Indian Ocean when compared to the Caribbean and Pacific. This study further supports the applicability of ITS2-DGGE in studies on Symbiodinium diversity, and highlights that potentially ecologically informative biogeographic patterns may be overlooked when only cladal designations are employed.     

Ecology of Patella species from the Cape peninsula,South Africa. IV. Desiccation

The importance of desiccation as of factor limiting zonation was investigated for 7 South African Patella Linnaeus species. Small specimens lost water fastest and were least tolerant to desiccation. Differences in shell shape largely explain differences in the rate of desiccation, both within and between species: tall conical shells with a small mouth circumference minimize water loss. Tolerance was closely related to zonation, but zonation was not correlated with either shell shape or rate of water loss. Two groups of limpets could be distinguished: migratory species which increase in tolerance as they age, correlated with the habit of moving up the shore; and non-migratory species which have a narrow range of tolerance. Zonation of the former group is probably limited by physical factors while the latter are behaviourally restricted to a narrow zone. The selective value of these strategies is discussed.     

The response of encrusting coralline algae to canopy loss: an independent test of predictions on an Antarctic coast

We assessed whether published observations of the ecology of encrusting coralline algae (Rhodophyta) from tropical and temperate coasts could be used to predict patterns and responses on a polar coast where such knowledge does not exist. On subtidal rocky coasts near Casey, East Antarctica, we detected a strong positive association of understorey encrusting coralline algae with canopies formed by the endemic alga Himantothallus grandifolius. The experimental removal of H. grandifolius caused corallines to bleach from red to pink/white concomitant with a decline in their photosynthetic activity. The magnitude of this decline (mean ± SE = 56.85±8.43%) was remarkably similar to that observed on temperate coasts (45.98±5.91%). Positive effects of nutrient enrichment of the surrounding water, hypothesized to alleviate the negative effects of canopy loss on encrusting corallines, were not detected. Removing H. grandifolius increased the intensity of photosynthetically active radiation and ultra-violet radiation reaching the substratum by three orders of magnitude, providing the basis for models invoking enhanced irradiance as the primary cause of the negative effects of canopy loss. Striking similarities among our results and those from tropical and temperate coasts suggest that responses of encrusting corallines to loss of canopies may have predictive properties across large distances and environmental gradients (tropical–temperate–polar).     

Climate heterogeneity modulates impact of warming on tropical insects

Evolutionary history and physiology mediate species responses to climate change. Tropical species that do not naturally experience high temperature variability have a narrow thermal tolerance compared to similar taxa at temperate latitudes and could therefore be most vulnerable to warming. However, the thermal adaptation of a species may also be influenced by spatial temperature variations over its geographical range. Spatial climate gradients, especially from topography, may also broaden thermal tolerance and therefore act to buffer warming impacts. Here we show that for low-seasonality environments, high spatial heterogeneity in temperature correlates significantly with greater warming tolerance in insects globally. Based on this relationship, we find that climate change projections of direct physiological impacts on insect fitness highlight the vulnerability of tropical lowland areas to future warming. Thus, in addition to seasonality, spatial heterogeneity may play a critical role in thermal adaptation and climate change impacts particularly in the tropics.     

Shell parameters in relation to zonation in mediterranean limpets

The relation of shell form to habitat in the Mediterranean limpets Patella caerulea (L.) and P. lusitanica (Gmelin) was investigated. P. caerulea shells were significantly longer, broader, but lower in height than P. lusitanica shells. The regression of the ratio (length plus breadth)/2 on the height of the shell was found to characterize the two limpet populations, with a small degree of overlap reflecting the distribution of the two limpets on the shore.     

Contrasting plant productivity-diversity relationships across latitude: the role of evolutionary history

The relationship between net primary productivity and biological diversity has been a central topic in ecology for several decades. The unimodal ("hump-back") relationship has been the most widely accepted for plants with the decrease in diversity at high productivity usually attributed to competitive exclusion. However, the relatively small species pool size under high productivity conditions may account for this pattern as well. Small species pool sizes for highly productive habitats are characteristic of temperate regions, where productive habitats for speciation and species migration have historically been rare. In contrast, productive habitats in the tropics have been relatively common during evolutionary history, resulting in large species pools. We hypothesize that evolutionary history contributes to the observed productivity-diversity relationship of plants, and that the productivity-diversity relationship differs between temperate and tropical regions. We investigated the productivity-diversity relationship patterns from 163 case studies throughout the world. Latitude described approximately 80% of the variation in the shape of the relationships. The unimodal relationship was found to dominate in the temperate zone, whereas the positive relationship was significantly more common in the tropics. We detected no influence due to methods of productivity measurement, but unimodal or positive productivity-diversity relationships were more likely within larger ranges of productivity. The length of the productivity gradient did not affect the latitudinal influence. In summary, the shape of the productivity-diversity relationship differs between temperate and tropical regions and the different evolutionary history of the local species pools is a probable cause for the difference.     

Ecological impacts on the limpet Lottia gigantea populations: human pressure over a broad scale on island and mainland intertidal zones

Here long-term monitoring data taken at 33 sites in southern and central California coast and islands were used to evaluate the size structure of the large intertidal limpet, Lottia gigantea in restricted-access and in easily accessible intertidal zones that encompass a wide range of ecological variables. Using multi-dimensional analysis of population size structures, we found that sites on islands and strictly protected mainland sites have significantly larger median limpet sizes and a greater range of limpet sizes than unprotected mainland sites, while no pattern occurs in latitudinal or regional comparison of sites. Although intertidal predators such as oystercatchers were not the primary focus of the monitoring efforts, extensive natural history notes taken during sampling visits support the argument that predation was not a primary cause for the size structure differences. Finally, substratum differences were determined not to have biased the observation of larger limpets in protected sites. In regard to human interactions with limpets, we conclude that the degree of enforcement against poaching is the better predictor of limpet size structure than proximity to population centers or visitation to intertidal sites.     

UV-radiation versus grazing pressure: long-term floating of kelp rafts (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) is facilitated by efficient photoacclimation but undermined by grazing losses

Large quantities of floating macroalgae are traveling in coastal waters of the SE Pacific and in other temperate climate zones. While afloat, these algae are potentially exposed to full solar radiation, including UVA and UVB, which can have profound effects on their physiological and growth performance. Latitudinal variations in UV-radiation (UVR) are hypothesized to affect floating algae differently with higher impacts at low latitudes than at high latitudes. In addition, UVR together with grazing might accelerate the demise of floating kelps. This hypothesis was tested with outdoor laboratory experiments in which sporophytes of the giant kelp Macrocystis pyrifera (L.) C. Agardh were exposed to a combination of different UVR regimes (PAR only, PAR + UV) and grazing at three sites along the Chilean coast (20°S, 30°S, and 40°S). A latitudinal trend in irradiance was detected with increasing values from 40°S to 20°S. Surprisingly, floating M. pyrifera responded with a high acclimation potential within this latitudinal UVR gradient. At 20°S, floating kelps were slightly sensitive to UVR, which was reflected in reduced blade growth. At 30°S, physiological responses were hardly affected by the prevailing irradiance but sporophyte growth and thus persistence mainly depended on the presence or absence of amphipod grazers. At high latitudes, grazing had only minor impacts on algal biomass and blade growth, and kelps thrived well under all tested environmental conditions. Overall, our results reveal that floating M. pyrifera was only slightly affected by UVR and that sporophytes can efficiently acclimate over a latitudinal UVR gradient that spans from 20°S to 40°S. Given this high acclimation potential, we suggest that these (and possibly other) positively buoyant algae are important dispersal agents over a wide range of temperate latitude conditions.     

Latitudinal variation in linear growth and other shell characteristics of Macoma balthica

Shells of the bivalve Macoma balthica (L.) were studied from locations throughout its geographic range on both sides of the North Atlantic. These observations were supplemented by records from the literature. North-south clines were detected in shell coloration, relative shell dimensions and shell growth parameters. Moreover, shells from the two sides of the Atlantic were so different (particularly in the south) that these populations can hardly be regarded as conspecific. It is, therefore, argued that latitudinal changes should be studied separately for the American and European populations. In America, growth rates increased continuously to the south, whereas in Europe maximal growth was observed at more intermediate latitudes.Virginia Institute of Marine Science Contribution No. 1251     

Assessment of factors affecting heart rate of the limpet Patella vulgata on the natural shore

 Heart rate variations of a population of the limpet Patella vulgata were monitored in the natural environment (Lough Hyne, southern Ireland) by non-invasive, optoelectronic recording. The heart rates of 145 limpets of different sizes, living on vertical and horizontal substrata, were measured both in air and water at different environmental temperatures, while the animals were inactive on their home scars. The heart rates of emersed, inactive limpets were positively related to air temperature and negatively related to limpet size. These relationships were similar for limpets on vertical and horizontal substrates. In contrast, no significant relationship between heart rate and temperature was found in submerged limpets, probably due to the narrow thermal range of the water during the study period. During submersion, a significant negative relationship between heart rate and size was evident for limpets on vertical surfaces but not for limpets on horizontal surfaces. In general, submerged limpets had a higher heart rate, 1.16 times that of limpets exposed to air. Moreover, the heart rates of nine animals were recorded while they were moving and while inactive on their home scars. Active limpets had a faster heart rate, 1.6 times that of limpets resting on their home scars. The dependence of heart rate on environmental temperature, size, respiratory medium and activity, as observed in limpets on the shore, agrees well with laboratory data and with previous findings of the correlation of oxygen consumption with the same factors. Such in situ measurements may, therefore, prove useful in attempts to determine natural levels of energy expenditure in models on the behaviour of foraging molluscs. Received: 5 September 1999 / Accepted: 17 April 2000     

Zoogeographical aspects of rocky-intertidal molluscan fauna of the Pacific coasts of Japan

Intertidal molluscan faunas were studied on the exposed rocky shores of the Pacific coasts of Japan (excluding Hokkaido) to determine distribution patterns and latitudinal gradients of taxa. Collections were made at spring low tide in 1980. Five types of distribution patterns were found: tropical, tropical-temperate, warm-temperate, temperate, and boreal. Many examples of parallel species displacement, the term which Thorson (1957) proposed and generalized, were found. Latitudinal gradients of molluscan taxon were observed. Frequency of the Archaeogastropoda in gastropods increases northward, while frequency of the Mesogastropoda and the Neogastropoda decreases. Frequency of predatory gastropods decreases northward, and that of herbivorous ones increases. Commonness indices show that the molluscan fauna of the study sites situated within the wide area between the Kanto and Kyushu are closely related with each other, and they can be regarded as one province. The Tohoku district is less related with this region, and the Okinawa district is the least related.