Phylogenetic Analysis of Conservation Priorities for Aquatic Mammals and Their Terrestrial Relatives, with a Comparison of Methods

BACKGROUND: Habitat loss and overexploitation are among the primary factors threatening populations of many mammal species. Recently, aquatic mammals have been highlighted as particularly vulnerable. Here we test (1) if aquatic mammals emerge as more phylogenetically urgent conservation priorities than their terrestrial relatives, and (2) if high priority species are receiving sufficient conservation effort. We also compare results among some phylogenetic conservation methods. METHODOLOGY/PRINCIPAL FINDINGS: A phylogenetic analysis of conservation priorities for all 620 species of Cetartiodactyla and Carnivora, including most aquatic mammals. Conservation priority ranking of aquatic versus terrestrial species is approximately proportional to their diversity. However, nearly all obligated freshwater cetartiodactylans are among the top conservation priority species. Further, ∼74% and 40% of fully aquatic cetartiodactylans and carnivores, respectively, are either threatened or data deficient, more so than their terrestrial relatives. Strikingly, only 3% of all 'high priority' species are thought to be stable. An overwhelming 97% of these species thus either show decreasing population trends (87%) or are insufficiently known (10%). Furthermore, a disproportional number of highly evolutionarily distinct species are experiencing population decline, thus, such species should be closely monitored even if not currently threatened. Comparison among methods reveals that exact species ranking differs considerably among methods, nevertheless, most top priority species consistently rank high under any method. While we here favor one approach, we also suggest that a consensus approach may be useful when methods disagree. CONCLUSIONS/SIGNIFICANCE: These results reinforce prior findings, suggesting there is an urgent need to gather basic conservation data for aquatic mammals, and special conservation focus is needed on those confined to freshwater. That evolutionarily distinct--and thus 'biodiverse'--species are faring relatively poorly is alarming and requires further study. Our results offer a detailed guide to phylogeny-based conservation prioritization for these two orders

Phylogenetic review of tonal sound production in whales in relation to sociality

Background It is widely held that in toothed whales, high frequency tonal sounds called \u27whistles\u27 evolved in association with \u27sociality\u27 because in delphinids they are used in a social context. Recently, whistles were hypothesized to be an evolutionary innovation of social dolphins (the \u27dolphin hypothesis\u27). However, both \u27whistles\u27 and \u27sociality\u27 are broad concepts each representing a conglomerate of characters. Many non-delphinids, whether solitary or social, produce tonal sounds that share most of the acoustic characteristics of delphinid whistles. Furthermore, hypotheses of character correlation are best tested in a phylogenetic context, which has hitherto not been done. Here we summarize data from over 300 studies on cetacean tonal sounds and social structure and phylogenetically test existing hypotheses on their co-evolution. Results Whistles are \u27complex\u27 tonal sounds of toothed whales that demark a more inclusive clade than the social dolphins. Whistles are also used by some riverine species that live in simple societies, and have been lost twice within the social delphinoids, all observations that are inconsistent with the dolphin hypothesis as stated. However, cetacean tonal sounds and sociality are intertwined: (1) increased tonal sound modulation significantly correlates with group size and social structure; (2) changes in tonal sound complexity are significantly concentrated on social branches. Also, duration and minimum frequency correlate as do group size and mean minimum frequency. Conclusion Studying the evolutionary correlation of broad concepts, rather than that of their component characters, is fraught with difficulty, while limits of available data restrict the detail in which component character correlations can be analyzed in this case. Our results support the hypothesis that sociality influences the evolution of tonal sound complexity. The level of social and whistle complexity are correlated, suggesting that complex tonal sounds play an important role in social communication. Minimum frequency is higher in species with large groups, and correlates negatively with duration, which may reflect the increased distances over which non-social species communicate. Our findings are generally stable across a range of alternative phylogenies. Our study points to key species where future studies would be particularly valuable for enriching our understanding of the interplay of acoustic communication and sociality

Limited role of character displacement in the coexistence of congeneric Anelosimus spiders in a Madagascan montane forest

Evolutionary and ecological theory predicts that closely related and similar species should coexist infrequently because speciation is more likely to occur allopatrically than sympatrically, and because co-occurring species with similar traits may compete for limited resources, leading to competitive exclusion or character displacement. Here we study the unusual coexistence of 10 similar congeneric species of Anelosimus spiders within a small forest fragment in Madagascar. We asked if these species radiated in sympatry or allopatry, and if there was evidence for local-scale character displacement in body size and other species-level traits. We sampled ∼ 350 colonies (6346 individuals) along a 2800 m transect. We identified colonies using morphology and DNA barcoding, and tested the monophyly of local and regional species assemblages with time-calibrated phylogenies. We used null model analysis and phylogenetic signal inference to test for patterns of segregation in body size, microhabitat, phenology, and seasonality of coexisting species. We found that all species belong to a Madagascan clade that radiated during the Pliocene, but that contemporary local assemblages are non-monophyletic. This is consistent with allopatric speciation during periods of global cooling and expansion of grasslands, and subsequent species assembly as forest fragments re-expanded and coalesced. We found no evidence for character displacement, except for overdispersion and even spacing in phenology: species were segregated by instars in a manner consistent with resource partitioning or maintenance of reproductive isolation. Overdispersion or even spacing in phenology may contribute to coexistence either through resource partitioning or mate recognition. However, there was no support for a scenario of resource partitioning and divergence of body size or other correlated morphological characters. These traits are better explained by evolutionary forces operating during speciation, rather than ecological forces operating during local community assembly

Webs in Vitro and in Vivo: Spiders Alter their Orb-Web Spinning Behavior in the Laboratory

Many studies of the elegant architectures of orb webs are conducted in controlled laboratory environments that remove environmental variability. The degree to which spider behavior in these circumstances resembles that of spiders in the wild is largely unknown. We compared web architecture and silk investment of furrowed orb weavers Larinioides corium\u27s (Clerck 1757) building webs in laboratory cages and spinning webs on fences in the field and found significant differences. The volume of major ampullate silk in radii was 53% lower in cage webs, primarily because the silk was 50% thinner, but also because spiders tended to spin 14% fewer radii than in fence webs. Cage spiders also invested about 40% less flagelliform silk and aggregate glue in the capture spiral, although the difference was not statistically significant, a trend primarily driven by a decrease in the length of the glue-coated capture spiral. These patterns were consistent with spiders reducing silk investment when building at new web sites while they assessed insect abundance. Differences in the type of substrate for web attachment, amount of available space, and condition may also have influenced web architecture. Cage webs were more symmetrical than fence webs, which displayed an unusual horizontal asymmetry that may have maximized their capture areas within the constraints of the available fence-railing attachment sites. Our findings suggest using caution when generalizing the properties of laboratory-spun webs to more natural conditions. More importantly, they demonstrate that orb spiders actively modify their behaviors when spinning webs under different conditions

Notes on the biology of Anelosimus pacificus Levi, 1963 (Theridiidae, Araneae)—evidence for an evolutionary reversal to a less social state

UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biologí

Elevation gradient affects the distribution and host utilisation of Zatypota anomala (Hymenoptera, Ichneumonidae) associated with mesh web weaving spiders (Araneae, Dictynidae)

The spatial distribution of parasitoids is closely linked to the distribution and ecological requirements of their hosts. Several studies have documented changes in the fauna composition of parasitoids in response to elevation, but data on parasitoids associated with spiders are missing. The koinobiont ichneumonid wasp Zatypota anomala is strictly specialised on spiders of the genus Dictyna (Dictynidae) in Europe. We examined the distribution of spiders of the family Dictynidae in forest ecotones in central Europe across a broad elevation gradient (110–1466 m a.s.l.). We checked the spiders for parasitism by Z. anomala. It was most abundant at the mid-elevations (median 712 m a.s.l., range 179–870 m a.s.l.). We identified four dictynid spider species as Z. anomala hosts. These were Dictyna arundinacea, Dictyna uncinata, Nigma flavescens, and Nigma walckenaeri. All four species and the genus Nigma were recorded as hosts for the first time. The parasitoids strongly preferred juvenile instars of their hosts. The body length differed between parasitised Dictyna and Nigma spiders (medians: 1.95 mm and 2.55 mm, respectively). The distribution of Dictyna and Nigma spiders overlapped along the elevation gradient, but parasitism incidence significantly differed between spider genera along the elevation gradient. Nigma was parasitized at lower elevations between 179–254 m a.s.l. and Dictyna at higher elevations between 361–870 m a.s.l. The phenology of Z. anomala is closely tied to the univoltine life strategy of its host spiders. The parasitoid female oviposits in autumn, and its offspring overwinter as larvae on the host, reach adulthood during spring, and pass the summer as an adult

Bioprospecting Finds the Toughest Biological Material: Extraordinary Silk from a Giant Riverine Orb Spider

Background Combining high strength and elasticity, spider silks are exceptionally tough, i.e., able to absorb massive kinetic energy before breaking. Spider silk is therefore a model polymer for development of high performance biomimetic fibers. There are over 41.000 described species of spiders, most spinning multiple types of silk. Thus we have available some 200.000+ unique silks that may cover an amazing breadth of material properties. To date, however, silks from only a few tens of species have been characterized, most chosen haphazardly as model organisms (Nephila) or simply from researchers' backyards. Are we limited to ‘blindly fishing’ in efforts to discover extraordinary silks? Or, could scientists use ecology to predict which species are likely to spin silks exhibiting exceptional performance properties? Methodology We examined the biomechanical properties of silk produced by the remarkable Malagasy ‘Darwin's bark spider’ (Caerostris darwini), which we predicted would produce exceptional silk based upon its amazing web. The spider constructs its giant orb web (up to 2.8 m2) suspended above streams, rivers, and lakes. It attaches the web to substrates on each riverbank by anchor threads as long as 25 meters. Dragline silk from both Caerostris webs and forcibly pulled silk, exhibits an extraordinary combination of high tensile strength and elasticity previously unknown for spider silk. The toughness of forcibly silked fibers averages 350 MJ/m3, with some samples reaching 520 MJ/m3. Thus, C. darwini silk is more than twice tougher than any previously described silk, and over 10 times better than Kevlar®. Caerostris capture spiral silk is similarly exceptionally tough. Conclusions Caerostris darwini produces the toughest known biomaterial. We hypothesize that this extraordinary toughness coevolved with the unusual ecology and web architecture of these spiders, decreasing the likelihood of bridgelines breaking and collapsing the web into the river. This hypothesis predicts that rapid change in material properties of silk co-occurred with ecological shifts within the genus, and can thus be tested by combining material science, behavioral observations, and phylogenetics. Our findings highlight the potential benefits of natural history–informed bioprospecting to discover silks, as well as other materials, with novel and exceptional properties to serve as models in biomimicry.Primary funding for this work came from the Slovenian Research Agency (grant Z1-9799-0618-07 to I. Agnarsson), the National Geographic Society (grant 8655-09 to the authors), and the National Science Foundation (grants DBI-0521261, DEB-0516038 and IOS-0745379 to T. Blackledge). Additional funding came from the European Community 6th Framework Programme (a Marie Curie International Reintegration Grant MIRG-CT-2005 036536 to M. Kuntner). The 2001 field work was supported by the Sallee Charitable Trust grant to I. Agnarsson and M. Kuntner and by a United States National Science Foundation grant (DEB-9712353) to G. Hormiga and J. A. Coddington. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.Peer reviewe

Phylogeography of a successful aerial disperser: the golden orb spider Nephila on Indian Ocean islands

Abstract Background The origin and diversification patterns of lineages across the Indian Ocean islands are varied due to the interplay of the complex geographic and geologic island histories, the varying dispersal abilities of biotas, and the proximity to major continental landmasses. Our aim was to reconstruct phylogeographic history of the giant orbweaving spider (Nephila) on western Indian Ocean islands (Madagascar, Mayotte, Réunion, Mauritius, Rodrigues), to test its origin and route of dispersal, and to examine the consequences of good dispersal abilities for colonization and diversification, in comparison with related spiders (Nephilengys) inhabiting the same islands, and with other organisms known for over water dispersal. We used mitochondrial (COI) and nuclear (ITS2) markers to examine phylogenetic and population genetic patterns in Nephila populations and species. We employed Bayesian and parsimony methods to reconstruct phylogenies and haplotype networks, respectively, and calculated genetic distances, fixation indices, and estimated clade ages under a relaxed clock model. Results Our results suggest an African origin of Madagascar Nephila inaurata populations via Cenozoic dispersal, and the colonization of the Mascarene islands from Madagascar. We find evidence of gene flow across Madagascar and Comoros. The Mascarene islands share a common 'ancestral' COI haplotype closely related to those found on Madagascar, but itself absent, or as yet unsampled, from Madagascar. Each island has one or more unique haplotypes related to the ancestral Mascarene haplotype. The Indian Ocean N. inaurata are genetically distinct from the African populations. Conclusions Nephila spiders colonized Madagascar from Africa about 2.5 (0.6-5.3) Ma. Our results are consistent with subsequent, recent and rapid, colonization of all three Mascarene islands. On each island, however, we detected unique haplotypes, consistent with a limited gene flow among the islands subsequent to colonization, a scenario that might be referred to as speciation in progress. However, due to relatively small sample sizes, we cannot rule out that we simply failed to collect Mascarene haplotypes on Madagascar, a scenario that might imply human mediated dispersal. Nonetheless, the former interpretation better fits the available data and results in a pattern similar to the related Nephilengys. Nephilengys, however, shows higher genetic divergences with diversification on more remote islands. That the better disperser of the two lineages, Nephila, has colonized more islands but failed to diversify, demonstrates how dispersal ability can shape both the patterns of colonization and formation of species across archipelagos.</p