Examination of the “null” model of connor and simberloff for species co-occurrences on Islands

Summary Among birds of the Bismarck Archipelago, only certain combinations of the species in a guild coexist on islands, and some species that are very similar ecologically have mutually exclusive distributions. Diamond (1975) interpreted these patterns as biologically significant, involving effects such as competition. Connor and Simberloff (1979) claimed such patterns to be not recognizably different from random, because they were scarcely distinguishable from those generated by a null distribution supposedly not incorporating competition.On examining the analysis by Connor and Simberloff, we find that it actually yields the opposite conclusion: three faunas tested have grossly non-random structures, while their test is unworkable for the fourth fauna. The method of Connor and Simberloff for generating a null distribution has many fatal weaknesses: dilution of relevant data from guilds with irrelevant data from the whole species pool; hidden incorporation of effects of competition into the constraints; inability to recognize a checkerboard, the extreme example of a distribution produced by competition; reliance on inefficient Monte Carlo simulations; severe constraints that sometimes make generation of a null distribution impossible; failure to weight species combinations; and failure to identify the direction of non-randomness or the species combinations most responsible.Finally, we use other recent studies by Simberloff and coleagues to examine the value of constructing null hypotheses in community ecology.     

Methods for detecting non-randomness in species co-occurrences: a contribution

Summary Comparison of co-occurrences between species on a group of islands with those expected from a randombased null model could provide evidence on community structure. However, it is difficult to decide on the appropriate null model. Gilpin and Diamond proposed a model and a test for departure from it, but this test is shown to indicate significant structure even when applied to a matrix of random numbers. An alternative method is suggested, using the distribution of Gilpin and Diamond's deviation as test statistic, but determining the expected distribution by Monto Carlo simulation, and using many such simulations as a randomisation test of significance. The null model used accepts the observed totals of occurrences for islands and species; it therefore offers a somewhat conservative test. Applied to the Vanuatu bird data that Gilpin and Diamond used, significant departure from a null model is seen, but with an excess of extreme negative associations, the opposite result from that given by Gilpin and Diamond's method. It is not possible to tell whether the negative associations are due to autecology, biogeography, or to interactions between species.     

The checkerboard score and species distributions

Summary There has been an ongoing controversy over how to decide whether the distribution of species is random — i.e., whether it is not greatly different from what it would be if species did not interact. We recently showed (Roberts and Stone (1990)) that in the case of the Vanuatu (formerly New Hebrides) avifauna, the number of islands shared by species pairs was incompatible with a random null hypothesis. However, it was difficult to determine the causes or direction of the community's exceptionality. In this paper, the latter problem is examined further. We use Diamond's (1975) notion of checkerboard distributions (originally developed as an indicator of competition) and construct a C-score statistic which quantifies checkerboardedness. This statistic is based on the way two species might colonise a pair of islands; whenever each species colonises a different island this adds 1 to the C-score. Following Connor and Simberloff (1979) we generate a control group of random colonisation patterns (matrices), and use the C-score to determine their checkerboard characteristics. As an alternative mode of enquiry, we make slight alterations to the observed data, repeating this process many times so as to obtain another control group. In both cases, when we compare the observed data for the Vanuatu avifauna and the Antillean bat communities with that given by their respective control group, we find that these communities have significantly large checkerboard distributions, making implausible the hypothesis that their species distributions are a product of random colonisation.     

Competitive exclusion,or species aggregation?

Summary There is a long-standing dispute over whether the analysis of species co-occurrence data, typically on islands in an archipelago, can disclose the forces at work in structuring a community. Here we present and utilise three scores S, C and T. S gives the mean number of islands shared by a species pair in the presence/absence data under study. The scores C and T are based on the way that a pair of species occurs on a pair of islands. When each species occurs on a different island, this adds to the checkerboard score C; if they occupy the same island, this increases the togetherness score T.In judging whether observed values of S, C and T are compatible with a null hypothesis assuming no species interaction, we follow Connor and Simberloff (1979) in generating a control group of (constrained) simulated incidence patterns.Presence/absence matrices can have paradoxical features, in combining a high mutual exclusion by species (checkerboardedness) with a degree of species aggregation that is also high. We show that this is in fact inevitable — that, given the usual contraints, C and T can differ only by a constant. This means that extreme checkerboardedness can be produced by forces making for species aggregation, just as well as by those making for avoidance.If we restrict our attention to a subset of species, the constraints are less rigid and the S, C and T scores are somewhat freer to vary. We consider the confamilial subsets in the Vanuatu archipelago as likely candidates for revealing any competition forces at work. Calculating the actual S, C and T scores for these subsets, we compare them with the corresponding scores in a sample of simulated colonization patterns.The actual species-distributions differ significantly from what we would expect if the colonization choices of different species were uncorrelated (save for some biological constraints). The confamilial species of the real world share more islands, and occur in a pattern less checkerboarded, and more aggregated, than their simulation counterparts. This suggests that competition pressures, if they exist, are overcome by countervailing factors.The method used is applicable in other ways, and to a wider class of problems, in analysing the forces behind community structure.     

Les environnements du Vanuatu de l’Holocène à nos jours : un état des lieux des connaissances

The first Vanuatu archaeological site discovery dates back to the 1960s, then the scientific and archaeological knowledge of this archipelago have increased. Nevertheless, the human activities–palaeoenvironmental changes relation in Vanuatu is still a matter of debate. In order to better determine this relation, reviewing the literature on past and actual environment is needed. This paper presents here a state of knowledge on Vanuatu geology, climate, biodiversity and archaeology, from the mid-Holocene to our present-day. The young archipelago results from strong volcanic and tectonic activity, due to the active subduction zone between Australian and Pacific plates. The atmosphere–ocean interactions determine the wet and dry seasons. Fauna and flora are principally derived from Southeast Asia. The occurrence of several species and subspecies endemic to the archipelago, if not to some islands, is probably due to the Vanuatu isolated location, the size of the islands and the climatic gradient between north and south. The first human populations reached the archipelago around 3200 yr BP. The Lapita culture, characterized by decorated potteries, is shared across the whole Vanuatu between 3200 and 2900 yr BP. After 2900 yr BP, cultures differ from one island to another. After 600 yr BP, the Polynesian culture dominates in Vanuatu.     

When snails inform about geology: Pliocene emergence of islands of Vanuatu indicated by a radiation of truncatelloidean freshwater gastropods (Caenogastropoda: Tateidae)

The South Pacific archipelago Vanuatu has a very complex geological history including three major phases of volcanism creating island belts and phases of repeated submergence and re‐emergence. An important issue for the evolution of the biota of Vanuatu ambiguously discussed in the geological literature is the question whether the entire archipelago has been submerged until the early Pleistocene or if at least parts of the island of Espiritu Santo have remained subaerial throughout the Pliocene. We used a time‐calibrated phylogenetic analysis of freshwater gastropods of the family Tateidae based on COI, 16S rRNA and ITS2 to infer the colonization history of Vanuatu. Our analyses suggested that Espiritu Santo was colonized c. 3 Mya. Espiritu Santo was probably the place of origin for the subsequent colonization of the island of Erromango (2 Mya) and the Pleistocene radiation across the remaining archipelago. We describe 10 new species largely based on morphological and anatomical data. The genetic data in particular of the species from the young islands are taxonomically incongruent probably due to incomplete lineage sorting typical for young radiations. In contrast, the paraphyly of Fluviopupa espiritusantoana appearing in three distant clades indicates either the existence of cryptic species or the long survival of the stem species of almost the entire radiation.     

Island area and species diversity in the southwest Pacific Ocean: is the lizard fauna of Vanuatu depauperate?

One island group suggested to be an exception to the species–area Relationship is the Vanuatu Archipelago, a group of 13 large and 80 small islands in the southwest Pacific Ocean. To test the hypothesis that the lizard fauna of the Vanuatu Archipelago does not meet the predictions of the species-area relationship, and thus is depauperate, we compare diversity among several island groups in the southwest Pacific: Fiji, the Loyalty Islands, New Caledonia, Samoa, the Solomon Islands, Tonga, and Vanuatu. We found that the lizard diversity of Vanuatu meets the pattern of diversity predicted by the species-area relationship. The Solomon Islands, the largest and least isolated oceanic archipelago considered, has the greatest species diversity and endemism of the oceanic islands. Inclusion or exclusion of island groups based on factors such as geologic history or faunal source affects the strength of the relationship between diversity, area, and history of emergence, and influences perceptions of diversity within individual archipelagos. In addition to island size, factors such evolutionary time scale, speciation, and archipelago complexity influence species richness on islands.     

The effect of insularity on the diversity of land birds in the Fiji islands: implications for refuge design

Ecologists have argued over rules of thumb that could be used to set priorities in configuring systems of reserves for preserving biological diversity. To evaluate these simple strategies, I assembled a particularly large and comprehensive data set on the land birds of the Fiji archipelago. I analyzed the species distribution on 220 islands to compare the running total of species preserved by differènt sequences of adding nature reserves to a hypothetical reserve system, treating each island as if it were a potential reserve. A strategy of maximizing the number of islands contributing to any given reserve area (maximum fragmentation) is much more effective at including species than a strategy of maximizing the size of the island components of a reserve (minimum fragmentation). Nevertheless the maximum fragmentation strategy is not a very good one. It is less effective than many random strategies, especially when about 2–10% of total area is to be set aside as reserve, and when only rare species are considered. A computer program was used to determine an optimal strategy by maximizing the number of additional species added for each unit of area added. This strategy is always substantially more effective at encompassing species diversity than either the maximum or minimum fragmentation strategies. It is suggested that the poor performance of the minimum fragmentation strategy is due principally to the presence of many smaller-island endemics within the Fiji archipelago. More generally, it is argued that the effect of fragmentation on species diversity depends on the geographic scale and isolation of the region under consideration. In these respects the Fiji Archipelago may be a particularly good model for continental reserve systems.     

The SLOSS dilemma: a butterfly case study

Butterfly species richness is examined on simulated archipelagoes of 2, 3, 4 and 5 holm oak forest fragments in the Guadalajara Province (central Spain). It is shown that there are more species on several small islands than on a single island. Also, species number increases with the number of fragments that form the archipelago, and with the average distance between islands within the archipelago. Thus, we conclude, at least for butterflies in a system of fragmented holm oak forests in central Iberia, that the best strategy in order to maximize the conservation of species richness is the creation of a net of some small and scattered reserves.     

The influence of gene flow and drift on genetic and phenotypic divergence in two species of Zosterops in Vanuatu

Colonization of an archipelago sets the stage for adaptive radiation. However, some archipelagos are home to spectacular radiations, while others have much lower levels of diversification. The amount of gene flow among allopatric populations is one factor proposed to contribute to this variation. In island colonizing birds, selection for reduced dispersal ability is predicted to produce changing patterns of regional population genetic structure as gene flow-dominated systems give way to drift-mediated divergence. If this transition is important in facilitating phenotypic divergence, levels of genetic and phenotypic divergence should be associated. We consider population genetic structure and phenotypic divergence among two co-distributed, congeneric (Genus: Zosterops) bird species inhabiting the Vanuatu archipelago. The more recent colonist, Z. lateralis, exhibits genetic patterns consistent with a strong influence of distance-mediated gene flow. However, complex patterns of asymmetrical gene flow indicate variation in dispersal ability or inclination among populations. The endemic species, Z. flavifrons, shows only a partial transition towards a drift-mediated system, despite a long evolutionary history on the archipelago. We find no strong evidence that gene flow constrains phenotypic divergence in either species, suggesting that levels of inter-island gene flow do not explain the absence of a radiation across this archipelago.     

Rissooidean freshwater gastropods from the Vanuatu archipelago

During expeditions to Santo and the Torres islands belonging to the Vanuatu archipelago in 2006 and 2007, ten new species of tateid gastropods confined to springs, the upper most, slowly flowing regions of streams or the groundwater had been discovered. These species were now described based on shell morphology and anatomy. In accordance with geography, these characters placed the species from Vanuatu between those from New Caledonia and Fiji, suggesting a stepping stone-like dispersal across the Pacific with an origin in New Zealand and the far end on the Austral islands. We also assessed the threat status of the new species according to the IUCN Red List criteria and concluded that they should be amended by explicit incorporation of the scale of potential human impact or stochastic natural events relative to the size of the habitat of organisms.     

Similarity and local co-existence of species in regional biotas

The notion of community-wide character displacement hypothesizes that locally co-existing sets of competing species should be less similar than expected when compared to random expectations from a broader regional species pool. Here I use a mechanistic approach to the niche concept to show how this expectation is dependent on the types of traits involved. I investigate how two different niche components, those that relate to species' requirements (or responses to environmental factors) versus those that relate to species' impacts (or effects on environmental factors), affect predictions about the similarity of locally co-existing species. In contrast with more conventional approaches that focus on species impacts, I focus on species responses to conclude that locally co-existing species should be more similar in such traits than expected on the basis of random assortment from a larger equilibrium regional biota. In addition, I explore the evolutionary implications of exceptions that might favour the co-existence of species with dissimilar traits (especially those that determine species' impacts on the environment) and conclude that these implications differ when species compete for shared resources, interact via shared predators, or interact via both mechanisms. The analysis developed in this paper shows that the co-existence of species that are more similar than expected by chance is not incompatible with the notion of strongly interacting species in saturated local communities near equilibrium.     

Endemic Seed Plants in the Bahamian Archipelago

The Bahamian archipelago consists of approximately 2,400 islands occurring in the Atlantic Ocean off the coasts of Florida, Cuba, and Hispaniola. In 1982 Donovan Correll and Helen Correll published the most current synopsis of the floristic diversity of this island chain. Their publication cited a total of 1,371 vascular plant species of which 114 seed plants were listed as endemic to the archipelago (~8 % of the native flora). In the last 30 years, additional herbarium collections and taxonomic studies have shown that a number of species previously indicated to be endemic to these islands also occur in other regions or have been taxonomically merged into other species. The current number of species considered endemic to the Bahamian archipelago is 89 (~6 % of the total flora). There are 50 endemic species that have a known distribution on one (31 species) or two island groupings (19 species). Biogeographical analyses of endemic plant distributions shows three distinct clusters of species: southern, central, and the northern islands, with a fourth cluster that includes islands with a small area and one medium size island that seems that has been underexplored (i.e., Little Inagua). We anticipate that understanding the conservation status of endemic species and their distributions will help to develop legislation to preserve this Bahamian natural heritage.     

Gentamicin resistance among salmonellae. A ten-year study, 1973–1982

A total of 8579 Salmonella strains received during 1973–1982 were tested for their antibiogram patterns against nine routinely used antibiotics including gentamicin. Of these, 380 strains (4.4%) showed resistance to gentamicin at levels of 10 g/ml and above. A high degree of resistance to gentamicin was recorded in 1979 (18.7%) and 1980 (9.4%). M.I.C. levels of strains received during 1982 were determined and it was found that some strains had levels as high as 160 g/ml. The comparative results of gentamicin resistance from 1973 to 1982 are presented and the public health significance of the alarming increase in two years (1979–1980) is discussed.     

Food-web models that generate constant predator-prey ratios

Summary An approximately constant ratio of number of predator species/number of prey species is observed in several natural communities, although the exact value of the ratio may vary with habitat and the types of organisms in the food web. We test the hypothesis that a constant predator/prey ratio can be generated by what Holt (1977) terms apparent competition and what Jeffries and Lawton (1984) call competition for enemy-free space. We create simple, two trophic-level communities by drawing species of predators and prey at random from a species pool, simulating their interactions using Lotka-Volterra models. The simulated food webs converge over successive periods of invasion and extinction to locally stable systems with approximately constant ratios of number of predator species/number of prey species, despite varying initial conditions. As expected, predator/prey ratios take different values depending upon the biology of the simulated species. We conclude that apparent competition between prey species via shared enemies may be one mechanism whereby approximately constant predator/prey ratios are generated in natural communities.     

The myth of constant predator: prey ratios

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated communities by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.     

Lichen species from the northeastern shore of Sørkapp Land (Svalbard)

In July and August 2005, the occurrence, distribution, and ecology of lichen species were studied in northeastern Sørkapp Land region (Spitsbergen) in areas undergoing the process of rapid glacier recession over the past century. A total of 48 lichen species were observed and two species (Acarospora heppii and Pycnothelia papillaria) are newly reported for the Svalbard archipelago.     

Invertebrate Invasions on Pacific Islands and the Replacement of Unique Native Faunas: A Synthesis of the Land and Freshwater Snails*Contribution no. 2001-001 of Bishop Museum's Pacific Biological Survey.

The once immense diversity of native Pacific island land snail species, with high single island or archipelago endemism, is declining dramatically. The native/endemic species are being replaced by a much smaller number of widespread tropical tramps, that is, those species that are most readily transported by humans. The 82 introduced (including 14 cryptogenic) land snail species recorded include some that were distributed accidentally by Pacific islanders before European exploration of the Pacific and that are now widespread. However, the majority are modern introductions, with many recent accidental introductions often associated with the horticultural trade. Native freshwater faunas were less diverse than the terrestrial faunas and exhibited much lower endemism. Among the 59 alien freshwater species recorded (including 38 cryptogenic species), the most diverse and widespread are the thiarids. Predation by and competition with these aliens (as well as habitat loss) are probably important mechanisms underlying the loss of native taxa, but almost no quantitative or experimental work has been done to demonstrate such ecological interactions. Prevention of further spread and of new introductions should be the main approach. Increased public education and development of public trust is essential to the success of these efforts.     

The Bahamian Flora: Plant Exploration,Living Collections,and DNA – A Journey Started in 1963

Dr. John Popenoe was Director of Fairchild Tropical Botanic Garden (FTBG) between 1963 and 1989. Dr. Popenoe was a strong supporter of the Bahamian flora, and in 1963 he obtained extramural funds to develop a living collection focusing on Bahamian plants that was established in FTBG. During his tenure FTBG supported the publication of the latest comprehensive flora of this archipelago. A project initiated by the first FTBG herbarium curator, William Gillis, but published by Donovan and Helen Correll (with illustrations of Priscilla Fawcett) in 1982. This living collection of Bahamian plants has supplied plant material for molecular phylogenetic studies worldwide. There are DNA phylogenies for only 10 of the 89 Bahamian endemics; most of the material of these phylogenies came from plants from these collections (three species) and/or from fragments of herbarium specimens collected by Donovan Correll (six species). Only two of the species included in these phylogenies are from collections unrelated to FTBG. Excluding species restricted to the Bahamas, material from 14 Caribbean Island endemics that are part of these collections has been used in phylogenetic studies. The available molecular phylogenies show that the Bahamian endemics are closely related to species from the West Indies or continental areas of the Caribbean Basin. There is also a paucity of population genetic studies based on DNA markers focusing on Bahamian plants. Only two of the four available population genetic studies are for a Bahamian endemic taxon.     

Formation of disjunct plant distributions in Northeast Asia: a case study of <Emphasis Type="Italic">Betula davurica</Emphasis> using a species distribution model

Repeated climate change during glacial and interglacial periods of the Quaternary led to mass migrations that resulted in disjunct distributions for many species. However, few studies have examined the processes that form disjunct distributions in Northeast Asia (NEA). In this study, we examined the disjunct distribution of Betula davurica Pall. in the Japanese archipelago. This species is a dominant canopy tree found in cool-temperate deciduous broad-leaved forests of continental NEA. We hypothesized that Quaternary climate change caused the present disjunct distribution pattern of this species. To test this hypothesis, we adopted a species distribution model and examined a series of potential habitats in the Last Glacial Maximum (LGM), the mid-Holocene, and the present. We generated models in MaxEnt with B. davurica presence as the response variable and six bioclimatic variables as predictor variables. During the LGM, projected potential habitats were distributed around the Korean Peninsula, East China, and the Japanese archipelago, excluding Hokkaido. In the mid-Holocene, habitats retreated both from East China and western Japan, remained unchanged in the Korean Peninsula and central Honshu mountains, and expanded to northern China, the Russian Far East, as well as northern Japan (Hokkaido). Thus, post-LGM global warming led to an expansion of B. davurica distribution to northern parts of continental NEA, along with a retreat in the Japanese archipelago. This shift in populations formed the present disjunct distribution.