Host utilization of the Brazil nut family (Lecythidaceae) by sympatric wood-boring species of Palame (Coleoptera, Cerambycidae, Lamiinae, Acanthocinini)

Estimates of the total number of arthropod species in existence are based, in part, upon assumptions about both the host specificity of tropical insects and their restriction to the forest canopy. It has been difficult to evaluate these estimates because of the paucity of available data. A newly discovered association between wood-boring beetles (Cerambycidae) and their host plants in the Brazil nut family (Lecythidaceae) inspired a year-long rearing project in the Neotropical rain forest of French Guiana. Branches severed from five species of Lecythidaceae yielded 1813 cerambycids belonging to 37 species. Three cerambycid species– Palame anceps (Bates), P. crassimana Bates, and P. mimetica Monné–accounted for almost half of the individuals reared. Each demonstrated a different pattern of host fidelity. Palame crassimana emerged from four of the five potential hosts, P. anceps emerged exclusively from a single host, and P. mimetica made a seasonal change in host affiliation. Although Palame spp. emerged from both ground level and canopy branches, they made a seasonal shift in stratum: they reproduced at both levels during the dry season, but exclusively at canopy level during the rainy season. Even specialized tropical insects may show greater flexibility in host utilization than some current hypotheses suggest.     

Uncovering tropical diversity: six sympatric cryptic species of Blepharoneura (Diptera: Tephritidae) in flowers of Gurania spinulosa (Cucurbitaceae) in eastern Ecuador

Diversification of phytophagous insects is often associated with changes in the use of host taxa and host parts. We focus on a group of newly discovered Neotropical tephritids in the genus Blepharoneura , and report the discovery of an extraordinary number of sympatric, morphologically cryptic species, all feeding as larvae on calyces of flowers of a single functionally dioecious and highly sexually dimorphic host species ( Gurania spinulosa ) in eastern Ecuador. Molecular analyses of the mitochondrial cytochrome oxidase-I gene from flies reared from flowers of G. spinulosa reveal six distinct haplotype groups that differ by 7.2–10.1% bp (uncorrected pairwise distances; N  = 624 bp). Haplotype groups correspond to six distinct and well-supported clades. Members of five clades specialize on the calyces of flowers of a particular sex: three clades comprise male flower specialists; two clades comprise female flower specialists; the sixth clade comprises generalists reared from male and female flowers. The six clades occupy significantly different morphological spaces defined by wing pigmentation patterns; however, diagnostic morphological characters were not discovered. Behavioural observations suggest specific courtship behaviours may play a role in maintaining reproductive isolation among sympatric species. Journal compilation  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 779–797. No claim to original US government works.     

Evolution and determinants of host specificity in the genus Lamellodiscus (Monogenea)

The evolution and determinants of host specificity in Lamellodiscus species (Monogenea, Diplectanidae) were investigated. The 20 known Mediterranean species were studied, all parasites of fishes from the family Sparidae (Teleostei). An index of specificity, which takes into account the phylogenetic relationships of their fish host species, was defined. The link between specificity and its potential determinants was investigated in a phylogenetic context using the method of independent contrasts. Host specificity in Lamellodiscus species appeared to be highly constrained by phylogeny, but also linked to host size. Mapping specificity onto the parasite phylogenetic tree suggests that specialist species do not represent an evolutionary dead end, and that specialization is not a derived condition. It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 431−443.     

Molecular genetic analysis of two taxa of the Pleurotus eryngii complex: P. eryngii (DC.Fr.) Quèl. var. eryngii and P. eryngii (DC.Fr.) Quèl. var. ferulae

With the use of isozymes and PCR-fingerprinting analysis molecular markers were found between the varieties Pleurotus eryngii var. eryngii and P. eryngii var. ferulae within the Pleurotus eryngii complex, which allowed the identification of the fruitbodies collected in Southern and Central Italy. The study of sympatric localities has shown that there is no gene exchange between them in the field. The post-mating barriers between these taxa are not yet completely efficient. However, in the field the gene pools of the two taxa appear isolated and associated with specific host plants: Eryngium campestre and Ferula communis . On the basis of the genetic and ecological differences observed and given the absence of gene exchange in sympatric localities, P. eryngii and P. ferulae are to be considered distinct biological species. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 125–136.     

Host selectivity,haustorial anatomy and impact of the invasive parasite Parentucellia viscosa on floodplain vegetative communities in Japan

The parasitic plant Parentucellia viscosa has been introduced recently to the Japanese floodplain. Because of its parasitic nature and high fecundity, P. viscosa may well become a major plant invader with a significant impact on floodplain ecosystems. Thus, a knowledge of the host selectivity of P. viscosa will not only provide basic biological information, but will also contribute to the understanding of floodplain vegetation conservation. We evaluated the host selectivity by comparing the observed numbers of haustoria with those expected from the relative below‐ground biomass. In addition, we examined the haustorial anatomy to determine whether haustoria are functional, and compared the above‐ground biomass of three functional plant groups (grasses, legumes and nonleguminous forbs) in intact and parasite removal quadrats. We found that haustoria were nonrandomly distributed amongst host species, suggesting host preference for Poaceae and Fabaceae. In addition, haustoria attached to certain forbs did not penetrate into the stele. The above‐ground biomass of grasses and legumes was increased significantly by the removal of P. viscosa, but the biomass of forbs did not change significantly. These results suggest that host preference depresses the performance of Poaceae and Fabaceae, thus affecting the competitive relationships among plants, meaning that P. viscosa may pose a serious threat to indigenous endangered legumes and grasses. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 69–78.     

The gall-inducing habit has evolved multiple times among the eriococcid scale insects (Sternorrhyncha: Coccoidea: Eriococcidae)

The habit of inducing plant galls has evolved multiple times among insects but most species diversity occurs in only a few groups, such as gall midges and gall wasps. This phylogenetic clustering may reflect adaptive radiations in insect groups in which the trait has evolved. Alternatively, multiple independent origins of galling may suggest a selective advantage to the habit. We use DNA sequence data to examine the origins of galling among the most speciose group of gall-inducing scale insects, the eriococcids. We determine that the galling habit has evolved multiple times, including four times in Australian taxa, suggesting that there has been a selective advantage to galling in Australia. Additionally, although most gall-inducing eriococcid species occur on Myrtaceae, we found that lineages feeding on Myrtaceae are no more likely to have evolved the galling habit than those feeding on other plant groups. However, most gall-inducing species-richness is clustered in only two clades ( Apiomorpha and Lachnodius  +  Opisthoscelis ), all of which occur exclusively on Eucalyptus s.s . The Eriococcidae and the large genus Eriococcus were determined to be non-monophyletic and each will require revision.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 441–452.     

Seasonality, microhabitat and cryptic variation in tropical salamander reproductive cycles

In many salamanders, most aspects of reproduction coincide with seasonal changes in abiotic factors such as temperature and humidity. However, while these reproductive patterns have been well documented for temperate salamanders, detailed data for tropical species are relatively sparse. I used histological techniques to examine temporal variation in the spermatogenetic cycles of four species of Guatemalan bolitoglossine salamanders ( Bolitoglossa occidentalis , B. rostrata , Dendrotriton bromeliacia and Pseudoeurycea goebeli ) from different microhabitats along an elevational gradient. All four species have mature sperm present in the testis throughout the year, irrespective of patterns of courtship and egg-laying. Additionally, the two species from more seasonal habitats ( B. occidentalis and B. rostrata ) exhibit significant but cryptic levels of variation in the amount of spermatozoa present in the testis not detectable by external appearance or the presence of secondary sexual characteristics. For the two species from less seasonal cloud forests ( D. bromeliacia and P. goebeli ), there were no detectable patterns to variation within the testis. Regional variation in climate undoubtedly influences reproductive cycles, however, microhabitat and the immediate environment are also important determinants of reproductive strategy.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 489–496.     

Specialization of rainforest canopy beetles to host trees and microhabitats: not all specialists are leaf‐feeding herbivores

Reliable estimates of host specificity in tropical rainforest beetles are central for an understanding of food web dynamics and biodiversity patterns. However, it is widely assumed that herbivores constitute the majority of host specific species, and that most herbivore species feed on leaves. We tested the generality of this assumption by comparing both plant host‐ and microhabitat‐specificity between beetle communities inhabiting the foliage (flush and mature), flowers, fruit, and suspended dead wood from 23 canopy plant species in a tropical rainforest in north Queensland, Australia. Independent of host tree identity, 76/77 of the most abundant beetle species (N ≥ 12 individuals) were aggregated on a particular microhabitat. Microhabitat specialization (measured by S_m and Lloyd's indices) was very high and did not differ between flower and foliage communities, suggesting that each newly‐sampled microhabitat has a large additive effect on total species richness. In accordance with previous studies, host specificity of foliage‐inhabiting beetles was most pronounced among herbivorous families (Curculionidae, Chrysomelidae). By contrast, host specificity among flower‐visitors was equally high among herbivorous and nonherbivorous families (e.g. Nitidulidae, Staphylinidae, Cleridae). Effective specialization (F_T) measures showed that traditional correction factors used to project total species richness in nonherbivorous groups fail to fully capture diversity in the flower‐visiting beetle fauna. These results demonstrate that host specialization is not concentrated within folivores as previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 215–228.     

Mechanisms driving the specificity of a myrmecophyte–ant association

In the understory of pristine Guianese forests, the myrmecophyte Hirtella physophora almost exclusively shelters colonies of the plant-ant Allomerus decemarticulatus in its leaf pouches. We experimentally tested three non-mutually exclusive hypotheses concerning phenomena that can determine the species specificity of this association throughout the foundation stage of the colonies: (1) interspecific competition results in the overwhelming presence of A. decemarticulatus queens or incipient colonies; (2) exclusion filters prevent other ant species from entering the leaf pouches; and (3) host-recognition influences the choice of founding queens, especially A. decemarticulatus . Neither interspecific competition, nor the purported exclusion filters that we examined play a major role in maintaining the specificity of this association. Unexpectedly, the plant trichomes lining the domatia appear to serve as construction material during claustral foundation rather than as a filter. Finally, A. decemarticulatus queens are able to identify their host plant from a distance through chemical and/or visual cues, which is rarely demonstrated in studies on obligatory ant–plant associations. We discuss the possibility that this specific host-recognition ability could participate in shaping a compartmentalized plant-ant community where direct competition between ant symbionts is limited.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 90–97.     

Floral scents in butterfly-pollinated plants: possible convergence in chemical composition

The study explores whether or not there are convergent patterns in floral scent composition among plant species that completely or partially rely on butterflies for pollination. Floral scent compounds were analysed from 22 flowering butterfly-pollinated plant species, representing 13 families which originate mainly from temperate North Europe but also from tropical and temperate America. Scents were collected using the dynamic headspace adsorption method and identified with coupled gas chromatography and mass spectrometry (GC-MS). In total, 217 floral scent compounds were identified, with the number per species ranging from 8 to 65. The major emerging pattern is the occurrence of certain compounds emitted exclusively by the flowers of many of the investigated species in major amounts – the benzenoids phenylacetaldehyde and 2-phenylethanol, the monoterpenes linalool and linalool oxide (furanoid) I and II and the irregular terpene oxoisophorone. It is likely that these compounds serve as a signal to attract pollinating butterflies, and may have evolved in conjunction with the sensory capabilities of butterflies as a specific group of pollinators. While there is convergence in terms of the compounds sharing this function there has been a geographical divergence in terms of their relative abundance. The predominance (in terms of both numbers and relative amount) of benzenoids in many of the scent blends of the European temperate species and of linalool and its derivatives in those of the American species constitute two discernible groups among these plants.  © 2002. The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 129–153.     

Pollen-host specificity and evolutionary patterns of host switching in a clade of specialist bees (Apoidea: Diadasia)

We examined the levels of pollen-host specificity in North American Diadasia (Hymenoptera: Apoidea), a clade of specialist bees. We analysed the scopal pollen loads of 409 individuals representing 25 of the 30 species of Diadasia that occur in North America. Each Diadasia species showed a preference for one of five plant families. However, the 25 species varied in their level of host specificity: the average percentage by volume of the preferred host in pollen loads ranged from > 99% to < 75%. In 17 of the 25 species, all or most individuals examined contained pure loads of one host taxon, while in eight species individuals were less specialized and carried mixtures of several unrelated host taxa. Mapping these host preferences onto a phylogenetic tree indicated that Malvaceae is the most likely ancestral host for the genus, and use of other hosts can be explained by a single switch to each of the other four host-plant families. Thus, most speciation events were not associated with a host switch; this pattern does not support host switching as a niche partitioning strategy to avoid competition. Diadasia species are more likely to use host-plant families that are used by other Diadasia and Emphorine bees; however, there was no evidence of residual adaptation to ancestral hosts. Diet breadth appears to be a labile trait: transitions from narrower to broader host use, as well as vice versa, were observed. The observed patterns of host-use evolution may be driven, in part, by host morphology and/or chemistry.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 487–505.     

How is host egg mimicry maintained in the cuckoo (Cuculus canorus)?

To investigate the evolutionary mechanism (host specificity vs. random searching) maintaining mimicry between cuckoo egg appearance and that of different European cuckoo Cuculus canorus hosts, we studied the level of mimicry between the appearance of C. canorus eggs and that of their hosts' eggs in different habitats in southern Finland by using ultraviolet-visible reflectance spectrophotometry. In the main habitat used by C. canorus for reproduction, eggs laid in nests of different host species differed in appearance. Host use by C. canorus was not related to the abundance of hosts, and the level of mimicry was not related to host abundance in the habitat. Furthermore, a close match between C. canorus egg appearance and that of host eggs within habitats was detected after removing the potentially confounding effect of host abundance. In the only two suitable host species nesting in trees (namely chaffinch Fringilla coelebs and brambling Fringilla montifringilla ) we detected changes in C. canorus egg appearance that paralleled those of the two host species. Thus our findings suggest the existence of a correlation between the appearance of C. canorus eggs and that of their hosts' eggs within different habitat types, and suggest that mimicry is maintained by strict host preferences by each C. canorus female when laying.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 57–68.     

Do canal‐cutting behaviours facilitate host‐range expansion by insect herbivores?

According to the escalation–radiation model of co-evolution, insect herbivores that acquire the ability to circumvent a plant defence enter a new adaptive zone and increase in species. How herbivore counter-adaptations to plant defences might lead to speciation is poorly understood. Studies of nymphalid butterflies suggest that the evolution of a broadened host range may be a critical step. This paper examines if leaf-feeding insects capable of deactivating defensive plant canals with canal cutting often have broad host ranges. A total of 94 species of canal-cutting insects were identified from the literature, including eight new canal cutters described in this paper. Only 27% of canal cutters with known host ranges are generalists that feed on plants in multiple families. The proportion of generalist canal cutters is similar or lower than estimates of generalists among phytophagous insects overall. Only five species, at most, of the canal-cutting generalists feed exclusively on plants with secretory canals. The paucity of generalists can be attributed in part to the considerable taxonomic distance separating canal-bearing plant families and to their corresponding chemical distinctiveness. The dependence of many canal-cutting species on host chemicals for defence would also favour specialization.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 715–731.     

The ecological differentiation of asexual lineages of cotton aphids: alate behaviour, sensory physiology, and differential host associations

The cotton aphid ( Aphis gossypii Glover) is a cosmopolitan pest species consisting of various host plant-associated lineages that appear to have different patterns of insecticide resistance, electrophoretic banding, and reproductive output across alternative host species. Debate continues as to whether these differences represent intraspecific variation or indicate a complex of cryptic or sibling species. We studied the host plant-related ecology of two asexual lineages of A. gossypii associated with the families Malvaceae and Cucurbitaceae, respectively, in particular the host plant finding and acceptance processes of the alate forms. The two lineages proved to be differentiated across host plants in terms of: (1) their landing and settling behaviour, nymphal production rate, and subsequent colony establishment (i.e. nymphal survival and development) and (2) their sensory abilities in relation to host plant-derived odours. Furthermore, genetic structuring according to host plant type was detected using the gene for elongation factor 1-α. The magnitude of the genetic differentiation across lineages was low, perhaps indicating a relatively recent divergence. The results demonstrate: (1) the presence of (at least) two cryptic species within the current definition of A. gossypii , as anticipated in previous studies, and (2) the practicalities of using a multidisciplinary approach to understanding the 'limits' of asexual species. If sexuals can be generated from each of these lineages, this conclusion will be open to further testing.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 503–519.     

How do host plant use and seasonal life cycle relate to insect body size: A case study on European geometrid moths (Lepidoptera: Geometridae)

We used European geometrid moths (>630 species) as a model group to investigate how life history traits linked to larval host plant use (i.e., diet breadth and host-plant growth form) and seasonal life cycle (i.e., voltinism, overwintering stage and caterpillar phenology) are related to adult body size in holometabolous insect herbivores. To do so, we applied phylogenetic comparative methods to account for shared evolutionary history among herbivore species. We further categorized larval diet breadth based on the phylogenetic structure of utilized host plant genera. Our results indicate that species associated with woody plants are, on average, larger than herb feeders and increase in size with increasing diet breadth. Obligatorily univoltine species are larger than multivoltine species, and attain larger sizes when their larvae occur exclusively in the early season. Furthermore, the adult body size is significantly smaller in species that overwinter in the pupal stage compared to those that overwinter as eggs or caterpillars. In summary, our results indicate that the ecological niche of holometabolous insect herbivores is strongly interrelated with body size at maturity.     

Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of molluscs at a New Caledonia site

Earlier studies in the tropical Indo-Pacific have grossly underestimated the richness of macrofauna species at spatial scales relevant to conservation and management as a result of insufficient collecting and sorting effort. A massive collecting effort involving 400 day-persons at 42 discrete stations on a 295-km² site on the west coast of New Caledonia, south-west Pacific, revealed 2738 species of marine molluscs. This is several times the number of species recorded from any area of comparable extension anywhere in the world. Spatial and habitat heterogeneity is high with 32% of the species collected at a single station. With 20% of the species represented by single specimens (0.4% of all catches), rare species make up a considerable proportion of the fauna. This justifies the parallel drawn between coral reefs and rain forests in terms of species diversity. The real richness of many soft-bodied marine taxa is probably underestimated, as evidenced by the fact that 28.5% of the mollusc species present at the study site are represented in the samples only by empty shells. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 421–436.     

Morphology and coexistence of congeneric ectoparasite species: reinforcement of reproductive isolation?

Assuming that differences or similarities in morphology among congeneric parasite species living in the same habitat are not a random pattern, several hypotheses explaining morphological differences were tested: (i) reproductive isolation, (ii) niche restriction resulting from competition, and (iii) niche specialization. Congeneric monogenean (platyhelminth) ectoparasites parasitizing the gills of one host species were used as an ecological model. Morphometric distances of the attachment organ and morphometric distances of the copulatory organ between species pairs were calculated, Levin's niche size and Renkonen niche overlap indices were applied. Our results support the prediction that the function of niche segregation is to achieve reproductive isolation of related species in order to prevent hybridization (reinforcement of reproductive barriers). Parasite species living in the same niche differ greatly in the size of copulatory organ. Moreover, species coexistence is facilitated by an increase in morphometric distances of copulatory organ and niche centre distances. Our results also show that species living in overlapping niches have similar attachment organs, which supports the prediction that morphologically similar species have the same ecological requirements within one host and suggests small effects of interspecific competition for the evolution of morphological diversity of attachment organs. Specialist adaptations also seem to facilitate species coexistence and affect the niche distribution within host species. Parasite species that can colonize more than one host species, i.e. generalists, occupy more distant niches within host species than strictly host-specific parasites. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76, 125–135.     

Molecular variation and speciation in Antitrichia curtipendula s.l. (Leucodontaceae, Bryophyta)

Molecular variation in Antitrichia curtipendula (Hedw.) Brid. s.l. was studied based on the nuclear internal transcribed spacer (ITS) and the chloroplast markers trn L- trn F and rpl 16, and analysed by neighbour joining (for ITS; recombination present), maximum parsimony (chloroplast markers) and TCS (haplotype network). Old World plus E North American populations belong to a different lineage than those of W North America. These are molecularly well differentiated and are treated as A. curtipendula and A. gigantea (Sull. & Lesq.) Kindb. Two distinct groups of Old World haplotypes are separated by one 'missing' haplotype and are interpreted as cryptic species. Tropical African populations share one ITS deletion and form a lineage within one of the cryptic species. Molecular variation within A. gigantea , within each of the two cryptic Old World plus E North American (except tropical African) haplotype groups, and among tropical African populations are of similar magnitude, suggesting that analogous mechanisms and similar time spans explain the found variation. Events related to Pleistocene climatic oscillations are suggested as having caused this differentiation within each group, whereas the African lineage was probably split off before this. Identical tRNA-Gly sequences were found in 33 specimens; new primers were designed for rpl 16 and ITS 1+2.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 341–354.     

Putative speciation events in Lamellodiscus (Monogenea: Diplectanidae) assessed by a morphometric approach

In the present study, we used morphometry as a proxy to study the microevolution of generalist Lamellodiscus (Monogenea, Diplectanidae) species, comprising gill parasites of sparid fish. We investigated 147 individuals, belonging to nine described species, regrouped in four morphotypes. Morphometric measurements were taken on sclerotized parts of the attachment organ. The formation of groups on the basis of the global morphometry within a host species, or between several host species, was assessed using both exploratory analyses (principal component analysis and clustering analysis) and statistical tests. We showed that: (1) for three out of four morphotypes, the global morphometry was significantly different according to host species used, and (2) the coexistence of two populations of Lamellodiscus elegans on Diplodus sargus could reflect an ongoing intra‐host speciation event. We suggest that generalist Lamellodiscus are undergoing specialization on their different hosts, which may lead to speciation. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 559–569.     

The spatial and temporal distributions of arthropods in forest canopies: uniting disparate patterns with hypotheses for specialisation

Arguably the majority of species on Earth utilise tropical rainforest canopies, and much progress has been made in describing arboreal assemblages, especially for arthropods. The most commonly described patterns for tropical rainforest insect communities are host specificity, spatial specialisation (predominantly vertical stratification), and temporal changes in abundance (seasonality and circadian rhythms). Here I review the recurrent results with respect to each of these patterns and discuss the evolutionary selective forces that have generated them in an attempt to unite these patterns in a holistic evolutionary framework. I propose that species can be quantified along a generalist–specialist scale not only with respect to host specificity, but also other spatial and temporal distribution patterns, where specialisation is a function of the extent of activity across space and time for particular species. When all of these distribution patterns are viewed through the paradigm of specialisation, hypotheses that have been proposed to explain the evolution of host specificity can also be applied to explain the generation and maintenance of other spatial and temporal distribution patterns. The main driver for most spatial and temporal distribution patterns is resource availability. Generally, the distribution of insects follows that of the resources they exploit, which are spatially stratified and vary temporally in availability. Physiological adaptations are primarily important for host specificity, where nutritional and chemical variation among host plants in particular, but also certain prey species and fungi, influence host range. Physiological tolerances of abiotic conditions are also important for explaining the spatial and temporal distributions of some insect species, especially in drier forest environments where desiccation is an ever‐present threat. However, it is likely that for most species in moist tropical rainforests, abiotic conditions are valuable indicators of resource availability, rather than physiologically limiting factors. Overall, each distribution pattern is influenced by the same evolutionary forces, but at differing intensities. Consequently, each pattern is linked and not mutually exclusive of the other distribution patterns. Most studies have examined each of these patterns in isolation. Future work should focus on examining the evolutionary drivers of these patterns in concert. Only then can the relative strength of resource availability and distribution, host defensive phenotypes, and biotic and abiotic interactions on insect distribution patterns be determined.