Ecological Expansion Research Articles

Expansion in geographical and morphological space drives continued lineage diversification in a global passerine radiation.

Why diversification rates vary so extensively across the tree of life remains an important yet unresolved issue in biology. Two prominent and potentially independent factors proposed to explain these trends reflect the capacity of lineages to expand into new areas of (i) geographical or (ii) ecological space. Here, we present the first global assessment of how diversification rates vary as a consequence of geographical and ecological expansion, studying these trends among 15 speciose passerine families (together approximately 750 species) using phylogenetic path analysis. We find that relative slowdowns in diversification rates characterize families that have accumulated large numbers of co-occurring species (at the 1° scale) within restricted geographical areas. Conversely, more constant diversification through time is prevalent among families in which species show limited range overlap. Relative co-occurrence is itself also a strong predictor of ecological divergence (here approximated by morphological divergence among species); however, once the relationship between co-occurrence and diversification rates have been accounted for, increased ecological divergence is an additional explanatory factor accounting for why some lineages continue to diversify towards the present. We conclude that opportunities for prolonged diversification are predominantly determined by continued geographical range expansion and to a lesser degree by ecological divergence among lineages.

Demosponge steroid biomarker 26-methylstigmastane provides evidence for Neoproterozoic animals.

Sterane biomarkers preserved in ancient sedimentary rocks hold promise for tracking the diversification and ecological expansion of eukaryotes. The earliest proposed animal biomarkers from demosponges (Demospongiae) are recorded in a sequence around 100 Myr long of Neoproterozoic-Cambrian marine sedimentary strata from the Huqf Supergroup, South Oman Salt Basin. This C30 sterane biomarker, informally known as 24-isopropylcholestane (24-ipc), possesses the same carbon skeleton as sterols found in some modern-day demosponges. However, this evidence is controversial because 24-ipc is not exclusive to demosponges since 24-ipc sterols are found in trace amounts in some pelagophyte algae. Here, we report a new fossil sterane biomarker that co-occurs with 24-ipc in a suite of late Neoproterozoic-Cambrian sedimentary rocks and oils, which possesses a rare hydrocarbon skeleton that is uniquely found within extant demosponge taxa. This sterane is informally designated as 26-methylstigmastane (26-mes), reflecting the very unusual methylation at the terminus of the steroid side chain. It is the first animal-specific sterane marker detected in the geological record that can be unambiguously linked to precursor sterols only reported from extant demosponges. These new findings strongly suggest that demosponges, and hence multicellular animals, were prominent in some late Neoproterozoic marine environments at least extending back to the Cryogenian period.

A giant dapediid from the Late Triassic of Switzerland and insights into neopterygian phylogeny

A new Triassic neopterygian is described on the basis of a large three-dimensional neurocranium from the Rhaetian (Late Triassic) of the Kössen Formation (Schesaplana, Grisons, Switzerland). CT scanning reveals neurocranial features similar to Dapedium, suggesting that this new genus, Scopulipiscis saxciput gen. et sp. nov., was deep-bodied and potentially durophagous, although no associated dental material is known. An expanded phylogenetic analysis of actinopterygians resolves Dapediidae as a clade (inclusive of Tetragonolepis), although fails to recover any characters supporting the monophyly of the genus Dapedium. Dapediids are resolved as stem holosteans, filling a conspicuous gap in early neopterygian relationships. Pycnodonts, previously suggested as either stem teleosts or the sister group to dapediids, are resolved as a clade on the neopterygian stem. Similarities between the new taxon described here and Dapedium provide insights into morphological disparity within early members of the group—suggesting that the ecological expansion of dapediids originated prior to the End-Triassic extinction—as well as contributing to a growing understanding of endocranial anatomy in Palaeozoic and Early Mesozoic actinopterygians.

A Trinitarian Basis for a “Theological Ecology” in Light of Laudato Si’

This article responds to Pope Francis’s call in Laudato Si’ for an ecological expansion of mission and seeks to provide it with theological support. This support comes by way of a trinitarian rendition of the missiological concept missio Dei. Drawing from Thomas Aquinas and Bernard Lonergan’s accounts of the trinitarian missions, it articulates a theological ecology (as opposed to an ecological theology), in which the traditional doctrine of God is the controlling motif. Through the missions of the Son and Holy Spirit, God transforms the moral-intellectual-volitional comportment of humanity and recruits them into a shared mission of environmental concern.

Ecological Expansion and Extinction in the Late Ediacaran: Weighing the Evidence for Environmental and Biotic Drivers.

The Ediacara Biota, Earth's earliest communities of complex, macroscopic, multicellular organisms, appeared during the late Ediacaran Period, just prior to the Cambrian Explosion. Ediacara fossil assemblages consist of exceptionally preserved soft-bodied forms of enigmatic morphology and affinity which nonetheless represent a critical stepping-stone in the evolution of complex animal ecosystems. The Ediacara Biota has historically been divided into three successive Assemblages-the Avalon, the White Sea, and the Nama. Although the oldest (Avalon) Assemblage documents the initial appearance of several groups of Ediacara taxa, the two younger (White Sea and Nama) Assemblages record a particularly striking suite of ecological innovations, including the appearance of diverse Ediacara body plans-in tandem with the rise of bilaterian animals-as well as the emergence of novel ecological strategies such as movement, sexual reproduction, biomineralization, and the development of dense, heterogeneous benthic communities. Many of these ecological innovations appear to be linked to adaptations to heterogeneous substrates and shallow and energetic marine settings. In spite of these innovations, the majority of Ediacara taxa disappear by the end of the Ediacaran, with interpretations for this disappearance historically ranging from the closing of preservational windows to environmentally or biotically mediated extinction. However, in spite of the unresolved affinity and eventual extinction of individual Ediacara taxa, these distinctive ecological strategies persist across the Ediacaran-Cambrian boundary and are characteristic of younger animal-dominated communities of the Phanerozoic. The late Ediacaran emergence of these strategies may, therefore, have facilitated subsequent radiations of the Cambrian. In this light, the Ediacaran and Cambrian Periods, although traditionally envisioned as separate worlds, are likely to have been part of an ecological and evolutionary continuum.

A hybrid land-water-environment model for identification of ecological effect and risk under uncertain meteorological precipitation in an agroforestry ecosystem

In this study, a hybrid land-water-environment (LWE) model is developed for identifying ecological effect and risk under uncertain precipitation in an agroforestry ecosystem. A simulation-based fuzzy-stochastic programming with risk analysis (SFSR) method is used into LWE model to reflect the meteorological impacts; meanwhile, it also can quantify artificial fuzziness (e.g., risk attitude of policymaker) and natural vagueness (e.g., ecological function) in decision-making. The developed LWE model with SFSR method is applied to a practical agroforestry ecosystem in China. Results of optimized planting scale, irrigative water schedule, pollution mitigation scheme, and system benefit under changed rainfall, precise risk-adoption and vague ecological function are obtained; meanwhile their corresponding ecological effects and risks are analyzed. It found that current LWE plans could generate massive water deficits (e.g., 23.22×106m3 in crop irrigation and 26.32×106m3 in forest protection at highest) due to over-cultivation and excessive pollution discharges (e.g., the highest excessive TP and TN discharges would reach 460.64 and 15.30×103 ton) due to irrational fertilization, which would increase regional ecological risks. In addition, fifteen scenarios associated with withdrawing cultivation and recovering forest based on regional environment heterogeneity (such as soil types) have been discussed to adjust current agriculture-environment policies. It found that, the excessive pollution discharges (TN and TP) could be reduced 12.95% and 18.32% at highest through ecological expansions, which would generate higher system benefits than that without withdrawing farmland and recovering forest. All above can facilitate local policymakers to modulate a comprehensive LWE with more sustainable and robust manners, achieving regional harmony between socio-economy and eco-environment.

Forest farming in Georgia, United States: Three potential crops

Abstract Forest farming is the cultivation of shade-tolerant crops under forest canopy. This agroforestry practice involves propagation, maintenance, harvesting, marketing, and overall economics of non-timber forest products, each of which falls into at least one category - medicinal herbs, foods, and/or ornamental plants. Interest in forest farming has been increasing, yet general information about this practice, including its advantages and disadvantages, are not readily available for the state of Georgia, United States. With declining indigenous plant species due to wild harvesting, landowners have the potential for economic and ecological expansion in this market. The goal of this review article is to evaluate the prospective economic outcomes of farming non-timber forest products. Basic background information, propagation methods and harvesting techniques are explained, and market values for recent years are provided for three non-timber forest products: black cohosh (Actaea racemosa) (medicinal herb), shiitake mushroom (Lentinula edodes) (food crop), and azalea (Rhododendron spp.) (ornamental plant). A list of other potential non-timber forest products in Georgia is also provided along with their uses and native status. The values of a few non-timber forest products of high demand, and consequently their potential profits, fluctuate widely from year to year leading to a wide range of reported profitability, depending on the source. Stability of the market depends on the steady availability of the product, which can be achieved once affected non-timber forest products are commonly cultivated instead of wild harvested.

Maternal care by Early Cretaceous cockroaches

Most cockroaches produce a special capsular structure, the ootheca, to enclose eggs for protection and maternal brood care. However, the origin and early evolution of the cockroach ootheca is poorly known, attributable to a lack of fossil evidence from the Mesozoic. Here, we report the earliest known cockroach fossil possessing an internally partitioned ootheca: Piniblattella yixianensis Gao, Shih & Ren sp. nov. is described from the mid Early Cretaceous Yixian Formation of China. The ootheca of this species measures 46–62% of its body length, contains 60–70 eggs, and exhibits the oviparity B reproduction mode, similar to that of the extant ectobiid and blattid cockroaches. In conjunction with several isolated ootheca specimens preserved in the mid-Cretaceous Myanmar amber (Burmite), we infer that the cockroach reproductive mode using the complete oothecate structure occurred during or before the mid Early Cretaceous. Maternal care associated with the oothecate condition in these Cretaceous cockroaches added a unique and major life-history trait, later resulting in true viviparity. This new life-history trait providing maternal protection and care may have been a key factor in the subsequent evolutionary diversification and ecological expansion of modern cockroach lineages.http://zoobank.org/urn:lsid:zoobank.org:pub:588C91F4-FB8D-43E5-8C2D-86D50393B851

Williamson Drive: Herbivory from a north-central Texas flora of latest Pennsylvanian age shows discrete component community structure, expansion of piercing and sucking, and plant counterdefenses

Insect herbivory during the Pennsylvanian–Permian transition interval (late Gzhelian) was estimated from the Williamson Drive locality of north-central Texas, USA. We investigated vegetational composition and the response of host plants to herbivores by characterization of insect-damage patterns. The bulk flora consists of four major plant groups: lycophytes, sphenophytes, pteridophytes, and pteridosperms, as well as unaffiliated morphotypes. Forty-six distinctive damage types (DTs) were recognized from 17 herbivorized plant-host taxa that were categorized into the five functional feeding groups (FFGs) of external foliage feeding (20 DTs), oviposition (11 DTs), piercing and sucking (9 DTs), galling (5 DTs), and wood boring (1 DT). The distributional patterns of FFGs and DTs on the most herbivorized plant hosts indicate several generalized and specialized patterns and organization into distinctive component communities. The overall herbivory level in the flora is elevated compared to other Cisuralian floras of north-central Texas, based on separate analyses of foliar surface area removed (0.86%) and frequency of herbivorized leaves (19.83%). One ubiquitous plant host, Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute and Zodrow, is disproportionately consumed, representing 35.75% of all specimens, but contributing 84.38% of the total herbivorized surface area, 60.87% of DT richness, and 55.44% of the DT abundance from all plant species. A notable feature is the abundance of nine piercing-and-sucking DTs on several hosts, representing three of the four new DTs described in this flora and indicating the ecological expansion of this FFG by Paleodictyopteroidea and earliest Hemipteroidea. There is evidence for physical and chemical antiherbivore defense afforded by trichomes and resin ducts of two major host plants, but the absence of seed predation is curious. Analyses of herbivory at Williamson Drive and four younger Cisuralian floras of the region support Feeny's apparency hypothesis. The overwhelming dominance of M. scheuchzeri herbivory at Williamson Drive, absent in earlier, peat-substrated Pennsylvanian floras, is typical of Cisuralian mineral-substrated plants.

JOHN BRADBY BLAKE, THE CHINESE TALLOW TREE AND THE INFRASTRUCTURE OF BOTANICAL EXPERIMENTATION

Recent studies of the spread of the invasive Chinese tallow tree, , Triadica sebifera (L.) Small, downplay the role of the dissemination of seeds in the 18th century compared with the impact of 20th century state‐sponsored agriculture. But the long‐term development of Chinese, then European and American botanical knowledge, even in cases like the tallow tree, can be shown to have enabled and legitimized ecological expansion and large scale environmental experimentation. John Bradby Blake, who obtained, experimented with, described and illustrated tallow tree seed in the 1770s, played an important role in the dissemination of tallow trees to the southeastern American colonies, the British Caribbean and northeast India.

Phylogeny and species diversity of Tasmanian mountain shrimps and their relatives (Crustacea, Anaspidesidae)

Anaspidacea is an enigmatic taxon within Malacostraca with uncertain phylogenetic affinities. One of the four families within Anaspidacea, the Anaspidesidae (representing in most respects the plesiomorphic conditions of the entire taxon), is endemic to Tasmania and shows a remarkable geographical and vertical distribution. Eleven species in three genera (Allanaspides, Anaspides and Paranaspides) are recognized to date, occurring in freshwater habitats such as lakes, rivers, streams, tarns and also in caves. We present the most detailed analyses to date of the phylogeny of Anaspidesidae including all eleven species based on molecular data (COI, 16S and 28S). In addition, the genetic differentiation within the morphologically recognized species is studied and a dating of speciation is given using a molecular clock approach (based on mt‐DNA only). A special focus is on the multiple instances of immigration into the subterranean habitat. Most of the morphologically delineated species are recognized by molecular data. Significantly, however, in at least two cases, the molecular data suggest more species than so far described, but further molecular sampling from a wider range of localities is required. Our molecular divergence estimates suggest the anaspidesids are younger than previously thought. Rather than having Eocene origins, our results suggest Early Miocene origins with primary diversification in Miocene and Pliocene. Entry into subterranean habitats appears to have occurred in the Pleistocene rather than Tertiary. Thus, rather than being a defensive response to rising temperatures in the Tertiary, entry into caves in the Pleistocene may represent opportunistic expansion into new habitats created by the action of glacial meltwaters on existing karst formations. As a result, environmental changes during the Pleistocene appear to have prompted ecological expansion of Anaspides by “opening” new subterranean habitats leading to increased diversification in the group.

Wood functional disparity lags behind taxonomic diversification in angiosperms

Early angiosperm fossil record shows a remarkably low wood structural diversity relative to that exhibited by reproductive structures. As wood anatomical traits are related to plant function, paucity of early wood structural differentiation suggests a delay of functional diversification relative to the explosive taxonomic accumulation of the group. We assembled a wood anatomy dataset of over 1000 extant species from Argentina, Brazil, China, Mexico, USA and Suriname to determine the disparification patterns of the cell types associated with the three main wood functional axes: conduction, support and storage. We found that most traits had a temporal shift in their evolutionary patterns, with early conservatism followed by high lability towards the present. Our results indicate that a surge in functional wood traits lability took place after the initial taxonomic diversification in the early Cretaceous and support the observed low wood structural diversity observed in the early angiosperm fossil record. Increased trait lability likely allowed evolutionary flexibility by lifting developmental constraints and boosted the evolution of different cell types involved in multiple wood functions and their associated ecological role. The diversification in wood function coincides with the escalation of angiosperm tree size and leaf hydraulic capacity and likely contributed to the subsequent ecological expansion of the angiosperms.

Coupled oceanic oxygenation and metazoan diversification during the early–middle Cambrian?

The early–middle Cambrian (Fortunian to Age 4) is characterized by a significant increase in metazoan diversification. Furthermore, this interval is marked by a prominent environmental and ecological expansion of arthropod- and echinoderm-rich biotas. Recent redox work has suggested that this shift occurred during stable or decreasing marine oxygen levels, suggesting that these paleobiological and paleoecological transformations were decoupled from a redox control. We tested this idea by conducting new paleoredox analyses on Age 2–Age 4 Cambrian outer shelf (Jiuqunao-Wangjiaping), slope (Wuhe-Geyi), and basinal (Zhalagou) sections of the South China Craton. Multiple sections indicate that mid-depth waters transitioned from anoxic conditions during Cambrian Age 2 to stable oxic conditions during Cambrian Age 4. These findings suggest a stepwise expansion of oxic waters from shallow to deep settings during the early–middle Cambrian, consistent with a redox control of metazoan diversification and ecological expansion. More broadly, despite the surge in redox work over the past decade, this study highlights the need for continued coupled redox and paleontological studies to directly test models about the links between the evolution of animals, ecosystems, and marine redox conditions.

Buchnera has changed flatmate but the repeated replacement of co-obligate symbionts is not associated with the ecological expansions of their aphid hosts.

Symbiotic associations with bacteria have facilitated important evolutionary transitions in insects and resulted in long-term obligate interactions. Recent evidence suggests that these associations are not always evolutionarily stable and that symbiont replacement, and/or supplementation of an obligate symbiosis by an additional bacterium, has occurred during the history of many insect groups. Yet, the factors favouring one symbiont over another in this evolutionary dynamic are not well understood; progress has been hindered by our incomplete understanding of the distribution of symbionts across phylogenetic and ecological contexts. While many aphids are engaged into an obligate symbiosis with a single Gammaproteobacterium, Buchnera aphidicola, in species of the Lachninae subfamily, this relationship has evolved into a 'ménage à trois', in which Buchnera is complemented by a cosymbiont, usually Serratia symbiotica. Using deep sequencing of 16S rRNA bacterial genes from 128 species of Cinara (the most diverse Lachninae genus), we reveal a highly dynamic dual symbiotic system in this aphid lineage. Most species host both Serratia and Buchnera but, in several clades, endosymbionts related to Sodalis, Erwinia or an unnamed member of the Enterobacteriaceae have replaced Serratia. Endosymbiont genome sequences from four aphid species confirm that these coresident symbionts fulfil essential metabolic functions not ensured by Buchnera. We further demonstrate through comparative phylogenetic analyses that cosymbiont replacement is not associated with the adaptation of aphids to new ecological conditions. We propose that symbiont succession was driven by factors intrinsic to the phenomenon of endosymbiosis, such as rapid genome deterioration or competitive interactions between bacteria with similar metabolic capabilities.

Geographical and ecological expansion in tropical butterflies of the genus Morpho in evolutionary time

Selected morphological. behavioral and ecological traits of Central and South American species of Morpho butterflies are examined as an attempt to account fot the present day geographical and ecological distributions of the genus in Central America

Evolution in an extreme environment: developmental biases and phenotypic integration in the adaptive radiation of antarctic notothenioids.

BackgroundOver the past 40 million years water temperatures have dramatically dropped in the Southern Ocean, which has led to the local extinction of most nearshore fish lineages. The evolution of antifreeze glycoproteins in notothenioids, however, enabled these ancestrally benthic fishes to survive and adapt as temperatures reached the freezing point of seawater (−1.86 °C). Antarctic notothenioids now represent the primary teleost lineage in the Southern Ocean and are of fundamental importance to the local ecosystem. The radiation of notothenioids has been fostered by the evolution of “secondary pelagicism”, the invasion of pelagic habitats, as the group diversified to fill newly available foraging niches in the water column. While elaborate craniofacial modifications have accompanied this adaptive radiation, little is known about how these morphological changes have contributed to the evolutionary success of notothenioids.ResultsWe used a 3D-morphometrics approach to investigate patterns of morphological variation in the craniofacial skeleton among notothenioids, and show that variation in head shape is best explained by divergent selection with respect to foraging niche. We document further an accelerated rate of morphological evolution in the icefish family Channichthyidae, and show that their rapid diversification was accompanied by the evolution of relatively high levels of morphological integration. Whereas most studies suggest that extensive integration should constrain phenotypic evolution, icefish stand out as a rare example of increased integration possibly facilitating evolutionary potential. Finally, we show that the unique feeding apparatus in notothenioids in general, and icefish in particular, can be traced to shifts in early developmental patterning mechanisms and ongoing growth of the pharyngeal skeleton.ConclusionOur work suggests that ecological opportunity is a major factor driving craniofacial variation in this group. Further, the observation that closely related lineages can differ dramatically in integration suggests that this trait can evolve quickly. We propose that the evolution of high levels of phenotypic integration in icefishes may be considered a key innovation that facilitated their morphological evolution and subsequent ecological expansion.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0704-2) contains supplementary material, which is available to authorized users.

Oceanic redox evolution across the end-Permian mass extinction at Shangsi, South China

The end-Permian mass extinction (EPME) was the most severe extinction event of the Phanerozoic. To investigate oceanic redox conditions around the EPME, we conducted a series of geochemical analyses, including iron speciation, trace element geochemistry, total organic carbon (TOC), and nitrogen isotopes of kerogen (δ15Nkero), around the EPME at the Shangsi section, South China. Three intervals with different redox conditions can be distinguished based on iron speciation. During the first interval (early and middle Changhsingian), bottom waters were pervasively euxinic. In the second interval (late Changhsingian and the EPME), bottom water fluctuated between oxic and anoxic, but non-sulfidic conditions, with the oxic mode becoming increasingly predominant over time. During the third interval, following the EPME, bottom waters appeared to have been consistently anoxic, although again not pervasively euxinic. This redox history is supported by enrichments in the authigenic fraction of redox-sensitive elements. These relatively high δ15Nkero values (>1.0‰) corresponding with Changhsingian euxinia suggest reducing bottom waters did not impinge on the photic zone. The shift to values near 0‰ within and above the EPME indicates an ecological expansion of diazotrophs. Stratigraphic trends of Nixs, Cdxs, Pxs, and TOC indicate that primary productivity was highest in the first interval, coinciding with euxinic bottom water, then decreased during the lower part of Interval 2 and remained at low level through the remainder of the studied section. The temporal relationship between redox condition and primary productivity suggests that euxinia was supported by high productivity and carbon export in the early Changhsingian. The expansion of N2 fixation coincides closely with the previously documented warming of the ocean surface at Shangsi and may be related to reduced oceanic circulation and suppressed vertical mixing. The temporal distribution of deepwater anoxia and euxinia suggests that they were not primary causes for the extinction of benthos at Shangsi.

Phylogeography and evolutionary history of the Crocidura olivieri complex (Mammalia, Soricomorpha): from a forest origin to broad ecological expansion across Africa.

BackgroundThis study aims to reconstruct the evolutionary history of African shrews referred to the Crocidura olivieri complex. We tested the respective role of forest retraction/expansion during the Pleistocene, rivers (allopatric models), ecological gradients (parapatric model) and anthropogenic factors in explaining the distribution and diversification within this species complex. We sequenced three mitochondrial and four nuclear markers from 565 specimens encompassing the known distribution of the complex, i.e. from Morocco to Egypt and south to Mozambique. We used Bayesian phylogenetic inference, genetic structure analyses and divergence time estimates to assess the phylogenetic relationships and evolutionary history of these animals.ResultsThe C. olivieri complex (currently composed of C. olivieri, C. fulvastra, C. viaria and C. goliath) can be segregated into eight principal geographical clades, most exhibiting parapatric distributions. A decrease in genetic diversity was observed between central and western African clades and a marked signal of population expansion was detected for a broadly distributed clade occurring across central and eastern Africa and portions of Egypt (clade IV). The main cladogenesis events occurred within the complex between 1.37 and 0.48 Ma. Crocidura olivieri sensu stricto appears polyphyletic and C. viaria and C. fulvastra were not found to be monophyletic.ConclusionsClimatic oscillations over the Pleistocene probably played a major role in shaping the genetic diversity within this species complex. Different factors can explain their diversification, including Pleistocene forest refuges, riverine barriers and differentiation along environmental gradients. The earliest postulated members of the complex originated in central/eastern Africa and the first radiations took place in rain forests of the Congo Basin. A dramatic shift in the ecological requirements in early members of the complex, in association with changing environments, took place sometime after 1.13 Ma. Some lineages then colonized a substantial portion of the African continent, including a variety of savannah and forest habitats. The low genetic divergence of certain populations, some in isolated localities, can be explained by their synanthropic habits. This study underlines the need to revise the taxonomy of the C. olivieri complex.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0344-y) contains supplementary material, which is available to authorized users.

Evolution of a complex behavior: the origin and initial diversification of foliar galling by Permian insects.

A central notion of the early evolution of insect galling is that this unique behavior was uncommon to rare before the diversification of angiosperms 135 to 125m.yr. ago. However, evidence accumulated during recent years shows that foliar galls were diverse and locally abundant as early as the Permian Period, 299 to 252m.yr. ago. In particular, a diversity of leaf galling during the Early Permian has recently been documented by the plant-damage record of foliar galls and, now, our interpretation of the body-fossil record of culprit insect gallers. Small size is a prerequisite for gallers. Wing-length measurements of Permian insects indicate that several small-bodied hemipteroid lineages originated early during the Permian, some descendant lineages of which gall the leaves of seed plants to the present day. The earliest foliar gallers likely were Protopsyllidiidae (Hemiptera) and Lophioneuridae (Thripida). Much of the Early Permian was a xeric interval, and modern galls are most common in dry, extra-tropical habitats such as scrubland and deserts. Plant-damage, insect body fossils, and the paleoclimate record collectively support the ecological expansion of foliar galling during the Early Permian and its continued expansion through the Late Permian.

Heritable symbiosis: The advantages and perils of an evolutionary rabbit hole

Many eukaryotes have obligate associations with microorganisms that are transmitted directly between generations. A model for heritable symbiosis is the association of aphids, a clade of sap-feeding insects, and Buchnera aphidicola, a gammaproteobacterium that colonized an aphid ancestor 150 million years ago and persists in almost all 5,000 aphid species. Symbiont acquisition enables evolutionary and ecological expansion; aphids are one of many insect groups that would not exist without heritable symbiosis. Receiving less attention are potential negative ramifications of symbiotic alliances. In the short run, symbionts impose metabolic costs. Over evolutionary time, hosts evolve dependence beyond the original benefits of the symbiosis. Symbiotic partners enter into an evolutionary spiral that leads to irreversible codependence and associated risks. Host adaptations to symbiosis (e.g., immune-system modification) may impose vulnerabilities. Symbiont genomes also continuously accumulate deleterious mutations, limiting their beneficial contributions and environmental tolerance. Finally, the fitness interests of obligate heritable symbionts are distinct from those of their hosts, leading to selfish tendencies. Thus, genes underlying the host–symbiont interface are predicted to follow a coevolutionary arms race, as observed for genes governing host–pathogen interactions. On the macroevolutionary scale, the rapid evolution of interacting symbiont and host genes is predicted to accelerate host speciation rates by generating genetic incompatibilities. However, degeneration of symbiont genomes may ultimately limit the ecological range of host species, potentially increasing extinction risk. Recent results for the aphid–Buchnera symbiosis and related systems illustrate that, whereas heritable symbiosis can expand ecological range and spur diversification, it also presents potential perils.