Geographic range limits of species

Understanding the forms that the geographic range limits of species take, their causes and their consequences are key issues in ecology and evolutionary biology. They are also topics on which understanding is advancing rapidly. This themed issue of Proc. R. Soc. B focuses on the wide variety of current research perspectives on the nature and determinants of the limits to geographic ranges. The contributions address important themes, including the roles and influences of dispersal limitation, species interactions and physiological limitation, the broad patterns in the structure of geographic ranges, and the fundamental question of why at some point species no longer evolve the ability to overcome the factors constraining their distributions and thus fail to continue to spread. In this introduction, these contributions are placed in the wider context of these broad themes.     

A macroevolutionary perspective on species range limits

Understanding the factors that determine the geographic range limits of species is important for many questions in ecology, evolution and conservation biology. These limits arise from complex interactions among ecology and dispersal ability of species and the physical environment, but many of the underlying traits can be conserved among related species and clades. Thus, the range limits of species are likely to be influenced by their macroevolutionary history. Using palaeontological and biogeographic data for marine bivalves, we find that the range limits of genera are significantly related to their constituent species richness, but the effects of age are weak and inconsistent. In addition, we find a significant phylogenetic signal in the range limits at both genus and family levels, although the strength of this effect shows interoceanic variation. This phylogenetic conservatism of range limits gives rise to an evolutionary pattern where wide-ranging lineages have clusters of species within the biogeographic provinces, with a few extending across major boundaries.     

Effects of habitat complexity on trophic interactions of three congeneric fish species

Aquatic Ecology - Habitat complexity can substantially alter trophic relationships, such as competitive and predatory interactions, between fish species. This study aimed to evaluate how trophic...     

Physiological tolerances account for range limits and abundance structure in an invasive slug

Despite the importance of understanding the mechanisms underlying range limits and abundance structure, few studies have sought to do so. Here we use a terrestrial slug species, Deroceras panormitanum, that has invaded a remote, largely predator-free, Southern Ocean island as a model system to do so. Across Marion Island, slug density does not conform to an abundant centre distribution. Rather, abundance structure is characterized by patches and gaps. These are associated with this desiccation-sensitive species'' preference for biotic and drainage line habitats that share few characteristics except for their high humidity below the vegetation surface. The coastal range margin has a threshold form, rapidly rising from zero to high density. Slugs do not occur where soil-exchangeable Na values are higher than 3000 mg kg⁻¹, and in laboratory experiments, survival is high below this value but negligible above it. Upper elevation range margins are a function of the inability of this species to survive temperatures below an absolute limit of −6.4°C, which is regularly exceeded at 200 m altitude, above which slug density declines to zero. However, the linear decline in density from the coastal peak is probably also a function of a decline in performance or time available for activity. This is probably associated with an altitudinal decline in mean annual soil temperature. These findings support previous predictions made regarding the form of density change when substrate or climatic factors set range limits.     

The energetics of starvation and growth after refeeding in juveniles of three cyprinid species

Synopsis Experiments were conducted to monitor changes in body mass and metabolic energy expenditure before, during, and after periods of starvation in juveniles of three species of cyprinids: Leuciscus cephalus, Chalcalburnus chalcoides mento, and Scardinius erythrophthalmus. During the starvation period all fish lost weight at about the same rate and the total amount of oxygen consumed during an experimental period of 20 h was about 40% lower in the starved than in the fed groups. Upon refeeding, both mass specific maintenance; and routine rates of metabolism as well as relative growth rates increased rapidly, the peaks of these increases being directly proportional to the length of the starvation period. Maximum compensatory growth was observed after four weeks of starvation in C. chalcoides and S. erythrophthalmus, with relative growth rates reaching 30% d^-1 during the first measuring interval after refeeding. The pattern of time-dependent compensatory growth displayed by these fish is similar to the responses of a colonial hydroid in which the rate of catch-up growth increased with the amount of stress to which the animals had been exposed. The exact cost of compensatory growth cannot be calculated because oxygen consumption and growth were not measured simultaneously. However, on the basis of data and calculations reported by Wieser & Medgyesy (1990) it appears that compensatory growth, if fuelled by the metabolic power indicated by our measurements of oxygen consumption, would have to be about twice as efficient as normal growth in the related species Rutilus rutilus.     

Physiological adaptations to entosymbiosis in three species of graffillid rhabdocoels

Three entosymbiotic graffillid rhabdocoels (Paravortex scrobiculariae, P. cardii and Graffilla buccinicola) from marine bivalve and gastropod molluscs show several physiological adaptations to their life-style which are intimately related to the nutritional physiology and ecology of their respective hosts.All three species feed on their hosts' partially digested food plus the cellular debris released at the end of the hosts' own digestive cycle. G. buccinicola supplements this diet by actively removing intact cells from the host's digestive epithelium. Host enzymes, ingested with the food, are utilized for digestion within the flatworms' gut; there is concomitant reduction in the types and amounts of endogenous enzymes and the gastrodermal gland cells characteristically found in free-living species are absent.Food reserves in the three species consist mainly of glycogen, following the pattern seen in other entosymbiotic flatworms (Turbellaria, Digenea, Cestoda); in P. scrobiculariae this primary adaptation, believed to be linked in all entosymbiotes to the ready availability of food and to high fecundity, probably has a secondary function, in relation to anaerobic respiration, of the type found in cestodes.Other adaptive features, closely correlated with host ecology, are the occurrence of a physiologically active haemoglobin in the brain and pharynx of P. scrobiculariae and, in this species and P. cardii, the differential occurrence and distribution of dehydrogenase systems concerned with aerobic respiration (Krebs cycle), the pentose phosphate shunt and anaerobic respiration (glycolysis).P. scrobiculariae and P. cardii are viviparous and the normal provision of yolk for embryonic nutrition is supplemented by direct passage of materials to the later larvae from the parental gut, thus facilitating extended development of the larva until its birth as an immature miniature adult.     

Patterns of foraging and range use by three species of neotropical primates

This paper describes the diet and range use patterns of the three species of primates in Santa Rosa National Park, Costa Rica (Ateles geoffroyi, Alouatta palliata, andCebus capucinus) and examines the variation in these variables as they relate to seasonal changes and concomitant changes in food availability. These three primate species were studied over a four-year period for a total of 24 months in the field. Santa Rosa National Park is in an area that experiences a long severe dry season in which little if any rain falls and the majority of the non-riparian trees lose their leaves. However, even though the three species were very flexible in terms of diet and range use, the behavioural variability did not correspond to changes in food availability or season.     

Phylogeny and species composition of five European species of Branchiobdella (Annelida: Clitellata: Branchiobdellida) reflect the biogeographic history of three endangered crayfish species

Branchiobdellid annelids are usually found as commensal symbionts associated with crayfish populations, but knowledge of their dispersion and ecology in Europe is generally scarce. We hypothesized that their geographic extension of species and populations may mirror the distribution history of their hosts. We analysed potential host specificities and the geographic distribution of species from the Italian and Austrian Tyrol and Carinthia by characterizing the morphological and genetic features. On the three indigenous crayfish species Astacus astacus, Austropotamobius pallipes and Austropotamobius torrentium , we identified four branchiobdellid species based on morphological characteristics: Branchiobdella hexodonta, Branchiobdella pentodonta, Branchiobdella balcanica and Branchiobdella parasita . In contrast to the morphological classification, phylogenetic analysis using mitochondrial cytochrome c oxidase I (CO-I) sequences identified five main lineages: B. balcanica, B. hexodonta, Branchiobdella italica, B. parasita and B. pentodonta . The arrangement of branchiobdellid species corresponded generally to the geographical distribution of their crayfish hosts' locations but also confirmed previous assumptions of crayfish translocations. Our study provides the first application of ideas on the association of freshwater crayfish and their ectosymbionts to be used for discussing the biogeography of crayfish populations. The phenotypical and genotypical analysis also demonstrated so far ignored effects of human activities at both macro-ecological and micro-ecological levels.     

Biogeography of the Amazon molly: ecological niche and range limits of an asexual hybrid species

Aim To understand the relative contributions of environmental factors, dispersal limitations and the presence of sperm donors in determining the distribution of the Amazon molly (Poecilia formosa), a sperm‐dependent unisexual fish species of hybrid origin. To explore niche similarities and/or differences between the hybrid and parental species. To evaluate whether large‐scale abiotic factors can explain a successful introduction of both P. formosa and Poecilia latipinna. Location South‐east United States, Mexico and Central America. Methods We used abiotic variables in ecological niche modelling (ENM) to identify regions with suitable conditions for the presence of the Amazon molly and its two parental species (P. latipinna and Poecilia mexicana). We also used a recently developed metric to calculate the degree of niche overlap between the hybrid and its parental species. Results ENM produced highly significant models [all area under the curve (AUC) > 0.99 for the three species]. Annual mean temperature and minimum temperature of the coldest month were the variables that best explained the distribution of the Amazon molly. With the exception of south Florida, few areas beyond the known distribution of the species were predicted to have suitable environmental conditions. The hybrid species niche overlaps partially with the parental species. However, given the available data, it is neither more similar nor more different than expected by chance. Main conclusions Two different processes are acting to limit the distribution of P. formosa. At the northern limit, although a sperm donor species is present further north, suitable environmental conditions are absent from nearby locations. At the southern limit, a sperm donor species is present and areas with good environmental conditions are present at nearby locations, suggesting that dispersal ability is the limiting factor. We found that the hybrid species overlaps in a similar way with both parental species while still having its own niche identity. This result may be explained by the fact that hybrid species inherit characteristics of two ecologically divergent species, which can result in intermediate or even transgressive phenotypes. These results support recent work on the role of hybridization in diversification.     

Inbreeding depresses short and long distance dispersal in three congeneric spiders

Dispersal is one of the most important precopulatory inbreeding avoidance mechanisms and subject to landscape related selection pressures. In small populations, inbreeding within and between populations may strongly affect population dynamics if it reduces fitness and gene‐flow. While inbreeding avoidance is generally considered to be a key evolutionary driver of dispersal, potential effects of inbreeding on the dispersal process, are poorly known. Here, I document how inbreeding within a population, so by mating among relatives, affects the survivorship and the dispersal behaviour of three congeneric spider Erigone species (Araneae: Linyphiidae) that differ in habitat preference and regional rarity. The three species were chosen as a model because they allow the assessment of both long and short distance dispersal motivation (respectively ballooning and rappelling) under laboratory conditions. Inbreeding reduced both long and short distance dispersal modes in the three congeneric species. Because survival was depressed after inbreeding, with a tendency of reduced survival loss in the rare and highly stenotopic species, energetic constraints are likely to be the underlying mechanism. Inbreeding consequently depresses silk‐related dispersal in three related spiders. This may induce an inbreeding depression vortex with important consequences for range expansion and metapopulation dynamics of aerially dispersing species from highly fragmented landscapes.     

Dietary and habitat niche partitioning in congeneric cryptobenthic reef fish species

Coral Reefs - Ecological niches hold critical information concerning the eco-evolutionary dynamics that govern biodiversity and abundance patterns. Cryptobenthic reef fishes account for...     

Consumer growth efficiencies: Their limits and relationships to ecological energetics

The relationship between biophage ecological efficiencies and growth efficiencies of individual animals is shown. The calculated theoretical maximum growth efficiencies are 82–88% for heterotherms and 53–56% for homeotherms. The factors that cause lowered observed growth efficiencies ($\text{x}\text{?}\text{= 53\%}$ for heterotherms) are discussed.     

Competition for S and N by associations of three annual range species in lysimeters

Summary Competition for S and N was studied on free draining lysimeters seeded to associations of three annual forages on Josephine loam (Typic Haploxerults), an important soil for revegetation to rangeland production. Forage yields increased and the botanical composition shifted toward subclover (Trifolium subterraneum L.) with added S. Sulfate uptake and plant S concentration were increased as a function of the S applied. Forage yields were related to plant S concentration and N:S ratios. Subclover and filaree (Erodium botrys (Cav.) Bertol.) obtained a larger proportion of their S from fertilizer-S than did soft chess (Bromus mollis L.). Total S uptake was largely from the applied fertilizer, 39% at the intermediate S level and 78% at high S. The replacement series design of the experiment and the use of labeled S and N permitted calculation of competition coefficients for fertilizer S, total S, soil N, and total N uptake for the species in pairwise associations. Sulfur was the factor limiting to subclover where S was not applied, and N was the limiting factor to soft chess and filaree in mixtures with subclover at high S where subclover was able to fix most of its own N. In the mixture of soft chess and filaree competition for S and N was about equally intense for both nutrients at all S levels, and filaree was dominant consistently. Comparison of two methods for estimating the fate of fertilizer S showed that differences in sulfur uptake and leachate losses over the controls provided significantly higher values for sulfur recovery than estimates based on the fate of³⁵S. Contribution of the University of California Hopland Field Station and Dept. of Agronomy and Range Science, Univ. of California, Davis, CA 95616.     

Impacts of global warming on Nezara viridula and its native congeneric species

Climate change and biological invasion are two of the most important ecological issues. Nezara viridula (SGS) is a good example of an alien species that increased in response to recent land use changes and global warming. The range limit of SGS coincided with the 5 °C isotherm of the mean monthly temperature for January. Since 2000, it has been warm enough for SGS to overwinter successfully outside its original range. Most invaded areas are now either occupied by SGS only or by mixed species of Nezara indicating that SGS is replacing its congeneric species, Nezara antennata, through interspecific mating.SGS population dynamics studies performed during 1961–1965 demonstrated that SGS abundance was density dependent and the independent processes worked alternately during the breeding season and winter. Although global warming would improve the winter survival of SGS adults, population density would be regulated at a new equilibrium specific to the area.Life tables demonstrated that all traits related to reproduction were poorly realized in the 2nd generation that occurs during summer. Heat stress syndrome due to global warming may be becoming apparent in SGS in spite of its subtropical origin. N. antennata may be able to avoid heat stress by aestivation. Whether SGS could continue to have a higher reproductive potential at the cost of heat stress over N. antennata is a significant ecological issue.     

Spatio-temporal distribution patterns of three stream-dwelling freshwater mussel species: towards a strategy for representative surveys

Conservation efforts for freshwater mussels (Unionoida) require a priori assessments of their populations’ status quo. Unfortunately, collection of representative census and other data for most European species is currently hampered by an insufficient understanding of inter- and intra-specific variation in distribution patterns. We assessed distribution and movement in three Unio crassus populations and one sympatric population of Anodonta anatina and Unio pictorum, respectively. We surveyed vertical movement across four sediment depths, and horizontal movement by mark-recapture technique in 3-month intervals. For all the populations, movement and the proportion inhabiting surface layers increased considerably from winter to spring/summer. Spatial aggregation levels remained stable for some populations, while others became increasingly randomly distributed during the study. One U. crassus population exhibited elevated mortality and displayed movement rates exceeding twice those of conspecific populations. The visible proportion of U. crassus populations differed by up to 69% between sites. Current monitoring guidelines in Europe often insufficiently account for the extensive inter- and intra-specific differences in spatio-temporal distribution patterns observed. We suggest developing internationally standardised protocols that specify sampling season and methodology. In particular, U. crassus surveys should be restricted to summer months, and hand-sampling is crucial for some populations.     

A comparison of the energetics of foraging of three species of Leptogenys (Hymenoptera, Formicidae)

Abstract. The three Leptogenys species L.nitida, L.schwabi and L.attenuata coexist in the coastal forests of South Africa and exhibit an array of foraging strategies ranging from individual foraging to group raiding. To determine whether there is a physiological basis for these strategies, the energetic cost of locomotion of individuals in these three species was determined. Carbon dioxide emission of voluntary running ants was measured using a flow-through technique, in order to determine their metabolic rate when running. The minimum cost of transport was constant over a range of temperatures (20–35°C), and similar for all three species (common value 212.96 ± 17.35 Jkg^_1m^_1). A comparison of the energy required to sustain representative foragers at 25°C indicated that it was energetically less expensive for L. nitida workers to forage than for the other two species. This may explain why L.nitida uses group raiding, while the other two species use individual foraging with limited recruitment for retrieval of large food items.     

Physiological energetics of the intertidal sea anemone Anthopleura elegantissima

Summary Lipid is the major energy storage molecule, and protein is highly conserved when food availability is low in high- and low-intertidal individuals of Anthopleura elegantissima. Under laboratory conditions zooxanthellae were lost from anemones, and tissue carbohydrate was mobilized to support metabolic demands. Routine catabolic requirements for carbohydrate are probably met by translocated photosynthate. Substrate-specific absorption efficiencies were greater than 95% for protein and lipid, but considerably lower (10 to 60%) for carbohydrate. This pattern of absorption is directly related to very low levels of dietary carbohydrate compared to the other substrates. High-intertidal anemones absorbed more lipid than low-intertidal anemones. Integrated budgets for organic nitrogen, the first reported for any cnidarian, indicate that low-intertidal anemones require larger daily rations than high-intertidal anemones to meet their higher metabolic demands for protein. These results are considered in relation to potential importance of zooxanthellae in nutrient cycling.     

Reliable and rapid discrimination of congeneric species by mtDNA SNP analysis by multiplex PCR: application on three Trachurus and two Mullus fish species as model cases

We present a reliable, time-saving, and cost-effective multiplex PCR assay for discriminating congeneric species. Three fish species of the genus Trachurus and two of the genus Mullus served as model cases. Our multiplex PCR method interrogates species-specific diagnostic mitochondrial single nucleotide polymorphisms (mtSNPs). We selected two sets of mtSNPs that are organized in two multiplex systems for the Trachurus and the Mullus species, respectively. In both systems, all individuals tested could be safely assigned to one of the three Trachurus or two Mullus species. This novel SNP typing system offers a convenient and robust DNA profiling system suitable for large-scale identification of commercial fish species, for species with cryptic larvae or during juvenile stages, as well as for wildlife forensics. Handling editor: Christian Sturmbauer