Shifts in Species Interactions Due to the Evolution of Functional Differences between Endemics and Non-Endemics: An Endemic Syndrome Hypothesis

Species ranges have been shifting since the Pleistocene, whereby fragmentation, isolation, and the subsequent reduction in gene flow have resulted in local adaptation of novel genotypes and the repeated evolution of endemic species. While there is a wide body of literature focused on understanding endemic species, very few studies empirically test whether or not the evolution of endemics results in unique function or ecological differences relative to their widespread congeners; in particular while controlling for environmental variation. Using a common garden composed of 15 Eucalyptus species within the subgenus Symphyomyrtus (9 endemic to Tasmania, 6 non-endemic), here we hypothesize and show that endemic species are functionally and ecologically different from non-endemics. Compared to non-endemics, endemic Eucalyptus species have a unique suite of functional plant traits that have extended effects on herbivores. We found that while endemics occupy many diverse habitats, they share similar functional traits potentially resulting in an endemic syndrome of traits. This study provides one of the first empirical datasets analyzing the functional differences between endemics and non-endemics in a common garden setting, and establishes a foundation for additional studies of endemic/non-endemic dynamics that will be essential for understanding global biodiversity in the midst of rapid species extinctions and range shifts as a consequence of global change.     

Co-occurrence of species with various geographical ranges, and correlation between area size and number of species in geographical scale

Co-occurrence of species of various geographical ranges is important to correct endemism evaluation. This co-occurrence is shown as non-hazardous. Influence of area size on species richness is assumed to be different with respect to endemic and non-endemic species. The territory of Israel and Sinai is subdivided into twenty biotic provinces. We segregated three hundred and twenty-five tenebrionid species inhabiting this territory into endemic, regional and ubiquitous species. Regression of the number of endemic species on the number of regional species is non-linear. Two distinct regression lines correspond to hot and cool areas. The number of ubiquitous species depends positively on numbers of both endemic and regional species, and negatively on their product. Ubiquitous species are predominantly synanthropic, and inability to tolerate competition with other tenebrionids is assumed as the basis of numerical relationships with other species. Correlation between numbers of endemic and non-endemic species of bird and mammal and size of area is analysed at the broad geographical scale. Relationships between area size evaluation and the numbers of endemic and non-endemic species are always different. The square root of the area km² is always more important in species richness determination than area itself. This variable is a linear characteristic of the area and its significance is discussed. Possible ecological interactions between species of various geographical ranges are also considered. A new method of evaluation of the level of faunal endemism is proposed.     

Invasion of pollination networks on oceanic islands: importance of invader complexes and endemic super generalists

Abstract. The structure of pollination networks is described for two oceanic islands, the Azorean Flores and the Mauritian Ile aux Aigrettes. At each island site, all interactions between endemic, non-endemic native and introduced plants and pollinators were mapped. Linkage level, i.e. number of species interactions per species, was significantly higher for endemic species than for non-endemic native and introduced species. Linkage levels of the two latter categories were similar. Nine types of interaction may be recognized among endemic, non-endemic native and introduced plants and pollinators. Similar types had similar frequencies in the two networks. Specifically, we looked for the presence of 'invader complexes' of mutualists, defined as groups of introduced species interacting more with each other than expected by chance and thus facilitating each other's establishment. On both islands, observed frequencies of interactions between native (endemic and non-endemic) and introduced pollinators and plants differed from random. Introduced pollinators and plants interacted less than expected by chance. Thus, the data did not support the existence of invader complexes. Instead, our study suggested that endemic super-generalist species, i.e. pollinators or plant species with a very wide pollination niche, include new invaders in their set of food plants or pollinators and thereby improve establishment success of the invaders. Reviewing other studies, super generalists seem to be a widespread island phenomenon, i.e. island pollination networks include one or a few species with a very high generalization level compared to co-occurring species. Low density of island species may lead to low interspecific competition, high abundance and ultimately wide niches and super generalization.     

Relationship between maximum leaf photosynthesis,nitrogen content and specific leaf area in balearic endemic and non-endemic mediterranean species

Gas exchange parameters, leaf nitrogen content and specific leaf area (SLA) were measured in situ on 73 C3 and five C4 plant species in Mallorca, west Mediterranean, to test whether species endemic to the Balearic Islands differed from widespread, non-endemic Mediterranean species and crops in their leaf traits and trait inter-relationships. Endemic species differed significantly from widespread species and crops in several parameters; in particular, photosynthetic capacity, on an area basis (A), was 20 % less in endemics than in non-endemics. Similar differences between endemics and non-endemics were found in parameters such as SLA and leaf nitrogen content per area (Na). Nevertheless, most of the observed differences were found only within the herbaceous deciduous species. These could be due to the fact that most of the non-endemic species within this group have adapted to ruderal areas, while none of the endemics occupies this kind of habitat. All the species-including the crops-showed a positive, highly significant correlation between photosynthetic capacity on a mass basis (Am), leaf nitrogen content on a mass basis (Nm) and SLA. However, endemic species had a lower Am for any given SLA and Nm. Hypotheses are presented to explain these differences, and their possible role in reducing the distribution of many endemic Balearic species is discussed.     

Links to rare climates do not translate into distinct traits for island endemics

Current models of island biogeography treat endemic and non-endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non-endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non-endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non-endemic species as functionally equivalent in island biogeography is not fundamentally wrong.     

Chance and necessity: land-snail faunas of Sao Miguel, Azores, compared with those of Madeira

The land-snail faunas of 23 sites from forest, disturbed andopen habitats were sampled on São Miguel, Azores archipelago.Collectively, the samples contained nearly all the known faunaof the island. Forest sites were the richest; while the numberof non-endemic species differed little between habitats, forestsheld more endemic species, many of which were widely distributed.Open habitats have no unique endemic species. These resultsare compared with similar data from Madeira. Although the twoislands are of the same size, and share the same Laurisilvaforest cover, their faunas differ in important aspects of diversity.While many non-endemic species are common to both islands, andoccur in similar numbers, endemic species are more numerouson Madeira, even though the numbers at individual forest sitesare, on average, fewer than on São Miguel. Many endemicsare confined to open, coastal habitats. These differences arediscussed in terms of the islands' histories, the range of habitatsavailable and the ease of dispersal from continental sources.Age, accidents of colonization and the range of natural habitatsavailable must have major effects on the faunas, but presentendemic faunal diversity is more a product of speciation andextinction than of immigration. (Received 13 May 2006; accepted 7 July 2006)     

Pattern and process in the ecological biogeography of European freshwater fish

1. Species lists for regions of Europe defined by Illies (1978, Limnofauna Europaea, 2nd edn. Gustav Fischer Verlag, Stuttgart), and augmented by information from Maitland (2000, Guide to Freshwater Fish of Britain and Europe. Hamlyn, London), are used to describe patterns in freshwater fish species richness and to examine the contribution of habitat preference, migration, body size and glacial history to these patterns. 2. The number of non-endemic species declines to the north and west, with increasing distance from the Ponto-Caspian region, the main source area, whereas endemic species richness declines only with latitude. 3. Habitat generalists tend to be migratory while riverine specialists are usually resident. Similar numbers of riverine species and generalists occur in Europe as a whole but generalists dominate in regional faunas and, to an increasing extent, in more isolated, formerly glaciated areas. Very few lacustrine specialists were found, reflecting the geologically ephemeral nature of lakes. Only 8% of riverine species have colonized glaciated areas, compared with more than half the generalist species, and the number declines rapidly with increasing distance from the source area. 4. Diadromous species show no geographical variation in species richness but potamodromous and resident species are affected by glaciation and by mountain and marine barriers. 5. The mean body size of regional faunas increases with latitude because there are relatively fewer small species in more distant, glaciated areas. 6. About half the species occurring in Europe are restricted to one region and the majority of these endemics occur in barrier regions with Mediterranean climates. Species in glaciated regions have much larger range sizes. Habitat preference and migration type, not body size, are the main determinants of range size. 7. Freshwater habitat availability varies across Europe with glaciated areas having more lakes of a given size than unglaciated areas. Catchment size is greatest at mid-latitudes. For a given catchment size rivers in glaciated areas are shorter. 8. The results support the notion that habitat variability, on both short and long time scales, favours colonization ability, which requires large body size. 9. As a result of their limited vagility northern fish faunas are depauperate, show high levels of plasticity and polymorphism and may show elevated speciation rates. The isolated southern faunas of the Iberian and Italian peninsulas and the Balkans are rich in endemic species but may be subject to extinctions because of the spread of the highly seasonal Mediterranean climate.     

Forest snail faunas from Crimea (Ukraine), an isolated and incomplete Pleistocene refugium

The land snail faunas of 26 forest sites and two open rocky sites in the Crimean Mountains were sampled in 2011. Of the 40 species found within the forests (about half the known fauna of Crimea as a whole), 28 were species with wide western Palaearctic distributions, and only eight were endemic to Crimea. While there were significant differences in the faunas of different sampling areas, these seemed to be a consequence of ecological differences among them rather than a product of geographical isolation and differentiation. Endemic species were large, and not entirely restricted to forest; known endemics not found in these forests are mainly typical of more open habitats. There is no local radiation of small species living in damp forest litter, as with Leiostyla species in the Transcaucasian forest refugium, and families such as the Clausiliidae with many endemic forest species in both Transcaucasia and the Carpathians are sparsely represented. The one endemic clausiliid genus, Mentissa, occurs in open as well as in wooded habitats. The present faunas are rather poor considering the soil conditions and climate, and the forests hold widespread species often associated with open habitats elsewhere. While there is evidence that these mountains provided a refuge for many animals and plants during glacial episodes further north, the forest snail fauna suggests that full forest cover did not survive throughout the Pleistocene. Rather, the present fauna contains endemics that survived in other habitats and widespread species with good powers of passive dispersal. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 424–433.     

Vicariance and endemism in a Neotropical savanna hotspot: distribution patterns of Cerrado squamate reptiles

Aim To test predictions of the vicariance model, to define basic biogeographical units for Cerrado squamates, and to discuss previous biogeographical hypotheses. Location Cerrado; South American savannas south of the Amazon, extending across central Brazil, with marginal areas in Bolivia and Paraguay and isolated relictual enclaves in adjacent regions. Methods We compiled species occurrence records via field sampling and revision of museum specimens and taxonomic literature. All species were mapped according to georeferenced locality records, and classified as (1) endemic or non‐endemic, (2) typical of plateaus or depressions, and (3) typical of open or forested habitats. We tested predictions of the vicariance model using biotic element analysis, searching for non‐random clusters of species ranges. Spatial congruence of biotic elements was compared with putative areas of endemism revealed by sympatric restricted‐range species. Effects of topographical and vegetational mosaics on distribution patterns were studied according to species composition in biotic elements and areas of endemism. Results We recorded 267 Cerrado squamates, of which 103 (39%) are endemics, including 20 amphisbaenians (61% endemism), 32 lizards (42%) and 51 snakes (32%). Distribution patterns corroborated predictions of the vicariance model, revealing groups of species with significantly clustered ranges. An analysis of endemic species recovered seven biotic elements, corroborating results including non‐endemics. Sympatric restricted‐range taxa delimited 10 putative areas of endemism, largely coincident with core areas of biotic elements detected with endemic taxa. Distribution patterns were associated with major topographical and vegetational divisions of the Cerrado. Endemism prevailed in open, elevated plateaus, whereas faunal interchange, mostly associated with forest habitats, was more common in peripheral depressions. Main conclusions Our results indicate that vicariant speciation has strongly shaped Cerrado squamate diversity, in contrast to earlier studies emphasizing faunal interchange and low endemism in the Cerrado vertebrate fauna. Levels of squamate endemism are higher than in any other Cerrado vertebrate group. The high number of recovered endemics revealed previously undetected areas of evolutionary relevance, indicating that biogeographical patterns in the Cerrado were poorly represented in previous analyses. Although still largely undocumented, effects of vicariant speciation may be prevalent in a large fraction of Cerrado and Neotropical biodiversity.     

Systematic review of the land snails of the Pitcairn Islands

The land snails (and semi-terrestrial molluscs) of the four islands that comprise the Pitcairn group are reviewed and the indigenous species illustrated. The strictly terrestrial molluscan faunas from the two atolls (Oeno and Ducie) are poor, like many other atolls in the Pacific. Each supports less than six species with wide geographical ranges. In contrast, the terrestrial molluscan fauna from Henderson Island, an uplifted atoll, is more diverse with at least 16 species belonging to seven families. Over half these taxa appear to be endemic, at least at the level of sub-species. Two species of semi-terrestrial molluscs have also been found on Henderson. Analyses of archaeological deposits in caves near the North Beach have revealed that at least a further six species of land snail formerly occurred on Henderson. The volcanic island of Pitcairn, the only island in the group still inhabited, supports the greatest number of terrestrial molluscs. Twenty-six species of land snail (and one semi-terrestrial pulmonate) were found living there during the recent expedition and a further three taxa were recognized amongst museum material. At least seven of these species are thought to be recent adventives and a further three are likely to have been prehistoric introductions. One Henderson (Georissa hendersoni) and three Pitcairn endemics (Pacificella filica, Sinployea pitcaimensis and Diastole tenuistriata) are formally described as new species. Some of the Pitcairn endemics occur in very restricted areas (less than a hectare) and it is important that measures should be taken to prevent the spread of invasive plants, such as rose-apple, that would threaten their survival.     

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.     

Modelling distribution patterns in a species-rich plant genus, Anthurium (Araceae), in Ecuador

Modelling potential species distributions has become a powerful tool for botanists in recent years. Using herbarium specimen data and GIS desktop software, we modelled the potential distribution of 36 endemic and 47 non‐endemic species of Anthurium (Araceae) in Ecuador based on mean annual temperature and humidity. Our results indicate the most important region for endemics in western Ecuador lies between the Andes and Coastal mountain ranges between 200 and 700 m, while for eastern Ecuador a belt of potential high diversity occurs directly along the foothills of the Andes under 1000 m. A very interesting result of this study highlights a site of predicted high species diversity at the borders of Guyas, Cañar, Bolivar, and Chimborazo, as well as sites within the Cordillera del Condor along the border with Peru. Potential richness for non‐endemic Anthurium species was similar to that of endemics with the inclusion of a large area of Amazonian lowlands in the east of the country. Over 40% of the protected areas in Ecuador occur in the eastern Amazonian lowlands, an area of low diversity for Anthurium endemics. Overall, for areas with potential high concentrations of endemic species identified in this study, only 3.1% are within Ecuador's protected areas.     

Larval supply is a good predictor of recruitment in endemic but not non-endemic fish populations at a high latitude coral reef

Despite extensive research, factors influencing the importance of pre- and post-settlement processes to recruitment variability remain ambiguous. Using a novel perspective, we investigated the potential influence of endemism on the relationship between larval supply and recruitment in reef fish populations at Lord Howe Island, Australia. Larval supply and recruitment were measured for three regional endemic and four widespread non-endemic species using light traps, artificial collectors, and underwater visual censuses. Recruitment was correlated with larval supply in endemics but not in non-endemics, likely due to a combination of low larval supply and post-settlement survival of non-endemics. Surveys also indicated that endemics were far more abundant and occurred in more locations than closely related non-endemics. These preliminary findings suggest that either local adaptation enhances recruitment in endemics through higher larval replenishment rates or reduced post-settlement mortality, populations of widespread species at the periphery of their range are poorly adapted to local environmental conditions and therefore experience lower and more variable settlement and post-settlement survival rates, or both.     

Diversity of mammals in the tropical–temperate Neotropics: hotspots on a regional scale

The tropical–temperate interface of the southern Neotropics harbours an interdigitating array of biomes (Puna, Monte, Chaco, Yungas). This topographic and climatically complex region needs urgent conservation efforts, as it is being transformed by human activities at an accelerating pace. We analyse georeferenced field records of mammal species in northwestern Argentina (provinces: Catamarca, Jujuy, Salta, Tucuman) in order to define biodiversity hotspots on the basis of 0.5°× 0.5° grid cells within northwestern Argentina according to total richness of mammal species, richness of megaspecies (species above 10 kg), and endemic species (species restricted to Argentina or neighbouring countries with shared biomes). The mammal fauna of northwestern Argentina is fairly well known (176 species). The biomes differ considerably in species richness (Puna low, Yungas high) and species composition. We found no significant difference between endemic and non-endemic species regarding cell occupancy or body size. Cell occupancy was not correlated to body size. Across grids, species richness, number of megaspecies as well as richness of endemics are all correlated to sampling effort. More than 50% of the species in the region are restricted to one or two biomes. Overall, the species turn-over between biomes in northwestern Argentina is high. Using a simple algorithm we identified 10 grid cells which covered 90% of the total number of recorded species, and contrast them with the protected areas. While the Puna and Yungas biomes are rather well protected, the arid and semiarid Monte and Chaco are in need of urgent attention in biodiversity conservation.     

Dispersal ability of island endemic plants: What can we learn using multiple dispersal traits?

Island endemic species are expected to have lower dispersal ability than their non-endemic congeners. Several studies have demonstrated differences in diaspore morphology between endemic species and their non-endemic congeners. It is, however, relatively difficult to translate the differences in morphology of the diaspores into differences in dispersal ability. To avoid this problem, we measured directly dispersal values (anemo-, hydro-, exozoo- and endozoo-chory) of 27 pairs of closely related endemic and non-endemic species from Canary Islands. We did not explicitly support the hypothesis about the loss of dispersal ability of island species. The comparison of pairs of endemic and non-endemic species showed the reduction in dispersal potential only for endozoochory. In many cases, endemic species had, in fact, the same or better dispersal ability than their non-endemic congeners. Higher dispersal ability of endemic species could have been evolved as a consequence of species subsequent dispersal to neighboring islands. As a support for this we found that the endemic species dispersing better than their non-endemic congeners also occupy more islands within the archipelago. We conclude that reduction of dispersal ability of species on islands may not be as general as previously expected and we need to take into account multiple species traits to understand the possible evolution of species dispersal potential.     

On the origin of the recent fauna of butterflies (Lepidoptera,Rhopalocera) of Vietnam

The following types of ranges of the Vietnamese butterflies are considered: 1) narrow ranges of endemic taxa, 2) continuous and mosaic ranges, 3) disjunct ranges, and 4) vicariant ranges. The endemic butterfly taxa mostly concentrate in isolated mountain areas of central Vietnam and show relationships with the Sino- Himalayan, Malayan, and local Indo-Burmese faunas. The ranges of most Vietnamese butterflies, both eury- and stenobiont species, combine the traits of continuous and mosaic distribution. Disjunctions in the ranges of Vietnamese butterflies vary from dozens to thousands kilometers. Similar habitats in different parts of Vietnam may have different though allied butterfly species. The origin of the recent butterfly fauna is hypothesized.     

Distribution and conservation significance of endemic species of flowering plants in Peru

Using the data published in the Catalogue of the Flowering Plants and Gymnosperms of Peru, we analyzed the elevational distributions of 5323 species reported as endemics from that country as a whole, for 10 families with the highest number of endemic taxa in Peru, and the distribution patterns of these species according to life form. We calculated the density of endemism (number of endemic species divided by area × 1000) and absolute number of endemic species among life forms and families, along an elevational gradient. Overall densities of endemics were 10–15 times higher at mid-elevation (2000–3500 m) than in the Amazonian lowlands (0–500 m). Absolute numbers of endemics peaked at 1500–3000 m for herbs, shrubs, and epiphytes, while trees, vines, and lianas showed maxima in the lowlands (0–500 m); yet densities of endemics for all life forms peaked at 1500–3000 m. Among the 10 families with the highest number of endemics, densities of endemics peaked at mid- to high elevation (1500–4500 m), but showed much disparity in the elevational distribution of absolute numbers of endemic species. Finally, the percentage of endemic species to total species is highest for herbs, shrubs, and epiphytes. Given that less than 10% of the land area for each of the montane zones (2000–4500 m) is protected compared to 13.5–29.9% in the lower elevations (0–1000 m), we recommend that priority be given to increasing the size of protected areas at mid- to high altitude in the Andean slopes to grant further protection in zones with the highest density of endemics. We also recommend that more emphasis be given to collecting and studying non-trees, since most endemic species belong to that class.     

Habitat selection and population trends in terrestrial bird species of Robinson Crusoe Island: habitat generalists versus forest specialists

Habitat loss and degradation on oceanic islands are key processes leading to population decline of endemic birds and facilitating the establishment of invasive bird species. In this study, carried out in the Robinson Crusoe Island, we assessed density and habitat selection of terrestrial bird species, including juan fernandez firecrown and juan fernandez tit-tyrant, two endemics, as well as green-backed firecrown and austral thrush, which apparently originate from the mainland. Results show that perturbed habitats contained a low density of the endemic species whereas the mainland species were significantly more abundant in perturbed scrub habitats. Bird species show different habitat selection patterns, with endemics selecting for native forest and mainland species selecting for perturbed habitats, or using them at random. Bird species experienced temporal trends in their overall population sizes, with the endemic tit-tyrant suffering a significant decline in its population size of about 63% between 1994 and 2009. Only mainland species exhibited temporal changes in habitat use, significantly reducing their densities in the preferred scrub habitats, possibly as a response to decreased habitat quality. Thrushes apparently were able to compensate the population decrease in one non native habitat type by using native forests, a habitat giving them the opportunity of preying on nests of endemic species. We conclude that endemic bird species behave as specialists whereas the mainland species must be treated as invasive generalists on Robinson Crusoe Island.     

Effects of Incubation Temperature on Seed Germination of Mediterranean Plants with Different Geographical Distribution Ranges

Little is known about consequences of the increasing global temperatures on seed germination and even less about the seed germination response of species with different geographical distribution ranges. In the present work, the effect of incubation temperature on seed germination of 31 species from the Iberian Peninsula is analyzed in relation to their geographical distribution range. We tested whether plants with different geographical distribution ranges (Iberian Peninsula endemics, Mediterranean, widely distributed species), respond similarly to incubation temperature using cross-species analysis as well as taking into account phylogenetic relationships among species. Seeds were incubated at 10°C, 15°C, 20°C and 25°C for 6?weeks in darkness. Distribution range appeared as a determinant of germination response to the temperature treatment independent of phylogeny. Widely distributed species were less sensitive than endemic or Mediterranean species to incubation temperatures, which may represent an advantage to colonize new areas or to tolerate changes in the environment. Although their ability to avoid germination during the warm and dry season might be favorable for endemic species in some situations, our results also suggest that under a future of temperature increase imposed by climate change, endemic plant species may be more vulnerable than widely distributed species.     

Biogeographical determinants for total and endemic species richness in a continental archipelago

We examined the relationship between plant species richness and biogeographical variables (island area, island maximum elevation, distance from nearest inhabited island, distance from nearest mainland) using a data set comprising 201 islands of the Aegean archipelago. We found that endemic species richness was strongly correlated to total species richness. Single-island endemic species richness was most strongly correlated to island maximum elevation, and then to island area, with an apparent small island effect for islands smaller than 47 km². Total species richness was most strongly correlated to island area (with no apparent small island effect), and less strongly correlated to island maximum elevation. Distance from the mainland or other inhabited islands displayed limited predictive value in our data set. The slope of the relationship between species richness and geographical factors (island area, elevation, distance from island/mainland) was steeper for endemic species richness than for total richness. Finally, the different scales of endemicity (single-island endemics, island group endemics and Aegean regional endemics) displayed similar qualitative trends and only differed quantitatively. Thus, we conclude that different biogeographical factors act as drivers for total species richness than for endemic species richness.