Areas, cradles and museums: the latitudinal gradient in species richness

Although numerous factors are postulated to be responsible for the gradient of increasing taxon richness towards lower latitudes, it has recently been suggested that the primary determinant is geographic area. This area model is appealing in its logic, but there is little empirical evidence to support it and several other mechanisms might also interact to obscure its effects. Nonetheless, the model has highlighted several fundamental issues concerning range size, speciation and extinction that, despite their considerable significance, remain poorly understood.     

Elevational patterns of species richness, range and body size for spiny frogs

Quantifying spatial patterns of species richness is a core problem in biodiversity theory. Spiny frogs of the subfamily Painae (Anura: Dicroglossidae) are widespread, but endemic to Asia. Using spiny frog distribution and body size data, and a digital elevation model data set we explored altitudinal patterns of spiny frog richness and quantified the effect of area on the richness pattern over a large altitudinal gradient from 0-5000 m a.s.l. We also tested two hypotheses: (i) the Rapoport's altitudinal effect is valid for the Painae, and (ii) Bergmann's clines are present in spiny frogs. The species richness of Painae across four different altitudinal band widths (100 m, 200 m, 300 m and 400 m) all showed hump-shaped patterns along altitudinal gradient. The altitudinal changes in species richness of the Paini and Quasipaini tribes further confirmed this finding, while the peak of Quasipaini species richness occurred at lower elevations than the maxima of Paini. The area did not explain a significant amount of variation in total, nor Paini species richness, but it did explain variation in Quasipaini. Five distinct groups across altitudinal gradient were found. Species altitudinal ranges did not expand with an increase in the midpoints of altitudinal ranges. A significant negative correlation between body size and elevation was exhibited. Our findings demonstrate that Rapoport's altitudinal rule is not a compulsory attribute of spiny frogs and also suggest that Bergmann's rule is not generally applicable to amphibians. The study highlights a need to explore the underlying mechanisms of species richness patterns, particularly for amphibians in macroecology.     

Sensitivity of intraspecific latitudinal clines of body size for tetrapods to sampling, latitude and body size

Recent studies have shown that most tetrapod groups (mammals,birds, chelonians, amphibians) show general intraspecific tendenciesfor increasing body size with latitude, whereas squamates (lizardsand snakes) show an intraspecific tendency towards decreasingbody size with latitude. Here I evaluate whether these sizetrends are general by using independent contrasts analysis toinvestigate the dependence of intraspecific size-latitude relationships(r), and the magnitude alone of size-latitude relationships([r]), for tetrapod vertebrates, on sample size, range of latitudessampled, average latitude sampled, and body size. Range of latitudessampled, average latitude sampled, and body size did not influencebody size-latitude relationships (r) or the magnitude aloneof body size-latitude relationship ([r]). Sample size did notinfluence size-latitude relationships (r), but did influencethe magnitude alone of size-latitude relationships ([r]), possiblyindicating increased precision of estimating size-latitude relationshipswith increased sampling. In short, intraspecific size-latituderelationships are similar for species of different sizes, occurringat different latitudes, sampled over different latitudinal ranges,and differing in number of populations sampled (though magnitudealone is influenced by sample size). These results suggest thatintraspecific size-latitude trends are general, and biologicallysignificant (i.e., are not artifacts of sampling), thus deservingexplanation.     

Interspecific patterns of species richness, geographic range size, and body size among New World venomous snakes

Many higher taxa exhibit latitudinal gradients in species richness, geographic range size, and body size. However, these variables are often interdependent, such that examinations of univariate or bivariate patterns alone may be misleading. Therefore, I examined latitudinal gradients in, and relationships between, species richness, geographic range size, and body size among 144 species of New World venomous snakes [families Elapidae (coral snakes) and Viperidae (pitvipers)]. Both lineages are monophyletic, collectively span 99° of latitude, and are extremely variable in body size and geographic range sizes. Coral snakes exhibit highest species richness near the equator, while pitviper species richness peaks in Central America. Species – range size distributions were strongly right-skewed for both families. There was little support for Bergmann's rule or Rapoport's rule for snakes of either family, as neither body size nor range size increased significantly with latitude. However, range area and median range latitude were positively correlated above 15° N, indicating a possible "Rapoport effect" at high northern latitudes. Geographic range size was positively associated with body size. Available continental area strongly influenced range size. Comparative (phylogenetically-based) analyses revealed that shared history is a poor predictor of range size variation within clades. Among vipers, trends in geographic range sizes may have been structured more by historical biogeography than by macroecological biotic factors.     

Predicting distributions of species richness and species size in regional floras: Applying the species pool hypothesis to the habitat templet model

The species pool hypothesis is applied here to the interpretation of ‘hump-shaped’ (unimodal) species richness patterns along gradients of both habitat fertility and disturbance level (the habitat templet). A ‘left-wall’ effect analogous to that proposed for the evolution of organismal complexity predicts a right-skewed unimodal distribution of historical habitat commonness on both gradients. According to the species pool hypothesis, therefore, the distribution of opportunity for net species accumulation (speciation minus extinction) should also have a corresponding unimodal central tendency on both habitat gradients. Two assumptions of this hypothesis are illustrated with particular reference to highly fertile, relatively undisturbed habitats: (i) such habitats have been relatively uncommon in space and time, thus providing relatively little historical opportunity for the origination of species with the traits necessary for effective competitive ability under these habitat conditions; and (ii) species that have evolved adaptation to these habitats are relatively large, thus imposing fundamental ‘packing’ limitations on the number of species that can ‘fit’ within such habitats. Based on these assumptions, the species pool hypothesis defines two associated predictions that are both supported by available data: (a) resident species richness will be relatively low in highly fertile, relatively undisturbed contemporary habitats; and (b) species sizes within regional floras should display as a right-skewed unimodal (log-normal) distribution. The latter is supported here by an analysis of data for 2,715 species in the vascular flora of northeastern North America.     

Latitudinal patterns in European ant assemblages: variation in species richness and body size

Using published distributions of 65 species from the British Isles and northern Europe, we show that ant assemblages change with latitude in two ways. First, as commonly found for many types of organisms, the number of ant species decreased significantly with increasing latitude. For Ireland and Great Britain, species richness also increased significantly with region area. Second, although rarely demonstrated for ectotherms, the body size of ant species, as measured by worker length, increased significantly with increasing latitude. We found that this body-size pattern existed in the subfamily Formicinae and, to a lesser extent, in the Myrmicinae, which together comprised 95% of the ant species in our study area. There was a trend for formicines to increase in size with latitude faster than myrmicines. We also show that the pattern of increasing body size was due primarily to the ranges of ant species shifting to higher latitudes as their body sizes increased, with larger formicines becoming less represented at southerly latitudes and larger myrmicines becoming more represented at northerly latitudes. We conclude by discussing five potential mechanisms for generating the observed body-size patterns: the heat-conservation hypothesis, two hypotheses concerning phylogenetic history, the migration-ability hypothesis, and the starvation-resistance hypothesis.     

Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.     

Ovarian egg size and number in relation to female size in five species of parasitoid wasps

Within five species of parasitoid wasps of the families Pteromalidae ( Morodora armata Gahan, Muscidifurax zaraptor K. & L., Nasonia vitripennis (Walker)), Bethylidae ( Goniozus legneri Gordh), and Tiphiidae ( Myzinum quinquecinctum (Fabricius)), we found significant positive correlations between female body size and both the number and the size of the mature oocytes present in the ovaries. These data are discussed with reference to current clutch size and egg size models and biological information available on these species.     

Effect of wasp size, physiological state, and prior host experience on host-searching behavior in a parasitoid wasp (Hymenoptera: Ichneumonidae)

Host-searching behavior in insects generally varies among individuals. A number of physiological and environmental factors can be involved in such individual variation. Here, a series of behavioral observations were made to highlight the importance of physiological state (i.e., number of mature eggs a female carries, amount of nutrient reserves, etc.) and learning state (i.e., prior host experience) on host-approaching behavior of parasitoids. Itoplectis naranyae (Hymenoptera: Ichneumonidae), a solitary endoparasitoid wasp attacking lepidopteran pupae and prepupae, was used as a test insect. The results show that female wasps with experience ovipositing on hosts 2 days before the test found hosts more quickly than did naïve wasps. Prior experience of host odor itself did not affect host-finding behavior, however. A single oviposition was enough for wasps to shorten time to find a host; additional experience had no significant effect on the efficiency of searching. The number of mature eggs a female carried had no effect on the time required to find a host regardless of prior host experience. The size of wasps, instead, was a significant factor when wasps had no prior host experience, and larger wasps found hosts more rapidly than did smaller wasps. Searching activity was not affected by how many hosts a female wasp had fed on before testing.     

Generic species richness and body mass in North American mammals: Support for the inverse relationship of body size and speciation rate

Generic species richness, the number of species per genus, is examined as a function of mean generic body mass for extant North American mammals. Species richness decreases as an inverse power function with increased mass, and the Spearman rank correlation coefficient of the logio transformed data is significant (r_s= ‐0.37). When the data are partitioned by trophic level, the relationship is not statistically significant for carnivores but strengthens for herbivores (r_s= ‐0.46). This interesting but incidental effect is due to the negligible number of diminutive and excessively large carnivores, which is in turn determined by foraging strategies. Alternate hypotheses for the “right‐skewed”; size distribution of modern North American mammals, such as disproportionate extinction of large species, differential species longevity, and a geographical scaling function, are rejected in favor of the proposition that elevated levels of speciation are restricted to animals of small body mass, as originally proposed by Gould and Eldredge (1977). This phenomenon is explained as a function of habitat restriction and particularly in herbivores, limited home range size. Aquatic mammals, regardless of body size, speciate rarely. Cope's Rule, the tendency of many animal groups to evolve towards large size, is understood as a probabilistic statement reflecting the phylogenetic tendencies of a disproportionately high number of small species alive at any given point in time.     

Loading constraints,body size and mating preference in Gammarus species

Size relationships among pairing Gammarus were examined with reference to two hypotheses for sexual size dimorphism and assortative mating among aquatic Peracarida (Crustacea). The sizes of pairing and non-pairing animals were compared in different experimental conditions where the size of one or both sexes was controlled. The experimental results present a complex picture which suggests that both sexual selection and loading constraints are likely to play a role in determining mating decisions in these animals.Department of Adult & Continuing Education, University of Durham     

The starvation-predation trade-off predicts trends in body size, muscularity, and adiposity between and within Taxa

The storage of lipids to buffer energy shortage may incur such costs as increased vulnerability to predation, and animals may be more muscular in order to reduce such costs. If muscle and lipid mass interact to determine survival, then both the muscularity and the adiposity of animals will be affected by factors such as predator density and food availability. Here we explore how adiposity and muscularity may depend on such factors. We confirm the expectation that adiposity should decrease with the risk of predation and increase with the frequency of interruptions to the food supply. More surprisingly, the predicted relationships between skeletal size, muscularity, and adiposity qualitatively depended on various factors: for example, adiposity should increase with foraging costs only for small animals and should decrease with total body mass if competition for food is intense. Furthermore, if the locomotive costs of carrying lipids are low, then adiposity should increase with body mass, whereas if such costs are high, then adiposity should decrease with body mass. These predictions are supported by observations of variation between and within species. Our approach demonstrates that broad patterns of body composition can be understood in terms of the fundamental ecological trade-off between starvation and predation.     

Relative influence of size,connectivity and disturbance history on plant species richness and assemblages in fragmented grasslands

Questions: What is the relative influence of size, connectivity and disturbance history on plant species richness and assemblages of fragmented grasslands? What is the contribution of small fragments to the conservation of native species pool of the region? Location: Tandilia's Range, Southern Pampa, Argentina. Methods: Cover of plants was registered within 24 fragments of tall‐tussock grassland remnants within an agricultural landscape using modified Whittaker nested sampling. We analysed the influence of site variables related to disturbance history (canopy height, litter thickness) and fragment variables (size, connectivity) on species richness (asymptotic species richness, slope of the species–area curve) as well as on species assemblages by multiple regressions analysis and canonical correspondence analyses, respectively. Cumulative area was used for analysing whether small fragments or large fragments are more important to species diversity in the landscape. Results: Asymptotic species richness was significantly influenced by site variables, in particular by Paspalum quadrifarium's canopy height, but not by fragment variables. Species assemblages were also affected by site variables (12.2% of total variation), but no additional portion of the species assemblage variability was significantly explained by fragment size and connectivity. Sampling of several small fragments rendered more exotic and native species than sampling of few large fragments of the same total area. Conclusions: Our results agree with previous studies reporting low sensitivity of species diversity to size and isolation of grassland fragments in fragmented landscapes and high sensitivity of species diversity to local variables. The higher capture of regional native species pool by small grassland fragments than by few larger ones of equivalent accumulated area highlights the value of small fragments for conservation.     

Mass, volume and dimensional distributions in biology, with special reference to cells

The positively skewed distribution of mass (weight) in biology was examined, and it was concluded all weight or masses can best be described by lognormal theory. Examples are given ranging in weight from viruses to humans. The accretion of mass proceeds as "more of the same" and primarily does not alter function. Immunologic properties must be carefully preserved. Small biologic entities can afford only small mass gains or losses while preserving their functionality; the larger the entity, the larger is the variability of mass functionally permitted. A well-established, convenient graphic method of analyzing a sample is described, with the advantages and pitfalls discussed. At the cellular level, volume and dimensions are lognormally distributed whenever the specific gravity (g/cu cm) is nearly homogenous among the particles or cells. In multicellular organisms, dimensions and forms (such as height and cranial circumference) are primarily the product of multifactorial genetic determinants and frequently appear as normal distributions. This paper discusses the meaning of the law of proportionate effects for small biologic objects, especially cells, and how an initial or "elementary" distribution may be conceived.     

Loudness of birdsong is related to the body size,syntax and phonology of passerine species

Songs of passerines are generally complex, long‐range acoustic signals, and are highly diverse across species. This diversity must nevertheless be shaped by the capabilities of the avian vocal physiology. For example, within species, loudness has been shown to trade‐off with aspects of song complexity. Here, I ask if such trade‐offs with loudness influenced the evolutionary diversification of song among passerines. Comparing perceived song loudness across > 140 European and North American species showed that loudness is positively related to body size and to singing with simple trilled syntax, and negatively related to aspects of syllable complexity. Syntax and syllable phonology together explained more variation than body size did, indicating that the acoustic design of songs is an important factor determining loudness. These results show for the first time that loudness covaries with, and possibly limits, song complexity across species, suggesting that a trade‐off with loudness shaped the evolutionary diversification of passerine song.     

Hierarchical large-scale to local-scale influence of abiotic factors in summer-fragmented Mediterranean rivers: structuring effects on fish distributions,assemblage composition and species richness

Summer rainfall shortage and low or no flows in Mediterranean rivers make it difficult for fish to attain their final preferenda and hinder the assembling of fish communities. We investigated in this study the degree to which large-scale features of the watersheds and reach-scale habitats correlated with a non-random structure of fish species' richness, composition and distribution under these conditions by intensively sampling a river system during summer and using Geographical Information Systems data. The results of multivariate ordination and association analyses showed that multi-scale factors were hierarchically related and that they correlated with non-random fish distributions and assemblage structure. They also highlighted large-scale factors shaping reach-scale water availability for fish during summer. Fish species' richness, composition and distribution varied along a gradient of water shortage and connectivity loss which were more severe with the smaller drainage area into the reaches upstream. Fish species' distributions along these gradients were strongly correlated with both life-history traits (adult size) and biogeographical origin (native vs. exotic). The results for the summer environmental segregation of exotic versus small endemic richness characteristics indicate strong species' interactions forcing the latter into the upstream, harsher seasonal habitats. Special fitness costs are to be expected in drought years, in regulated rivers, and in general, in a climate change scenario in Mediterranean river networks. The study emphasizes that the conservation of the threatened Mediterranean freshwater fish faunas requires operating at the proper spatial scale. In particular, reach-scale habitat improvements are adequate but not enough during the summer in fragmenting river networks, unless there is proper watershed-scale management of the flow regimes.     

Species richness of saproxylic beetles in woodlands is affected by dispersion ability of species,age and stand size

This study examines the occurrence of saproxylic beetles in woodlands of different size and age and their potential to leave woodland areas and cross open grassland in northern Germany. The beetles were recorded by emergence traps and flight-intercept-traps. The investigated sites were dominated by beech, oak and alder, but other tree species also occurred in low abundance. Species richness showed a positive relation to stand size and age of woods. Both total and rarefaction species richness were the lowest in small and young woods and highest in old and large woods. Species richness decreased asymptotically from the inner-wood habitat to a distance greater than 80 m from the wood margin. 80 species were classified into 46 low mobile species found at a distance <30 m from wood margins and 34 high mobile species found >30 m from wood margins. The most mobile species were found the most frequently in all woods; but they contributed less to species richness in wood stands than did the species with low mobility. The contribution of the least mobile species to species richness in wood stands increased with the age and size of the stands, with the effect of stand size being the greater. We conclude that in our study region woods larger than 100 ha are necessary to maintain the highest richness of the least mobile saproxylic beetles.