Differences In Adult Body Size Research Articles

Seasonality, body size, and maturation time in the neotropical grasshopper Sphenarium histrio across an altitudinal gradient

AbstractIn insects, male mating success and female fecundity usually increase with body size. However, natural selection favors faster maturation, reducing the risk of pre‐reproductive death, when the reproductive season is short in habitats located at high altitudes or far from the equator. Also, if males that mature earlier than females under these conditions increase their mating opportunities, protandry may evolve in their populations. Nonetheless, as body size is strongly correlated with maturation time in insects, faster sexual maturation is reached at the expense of having a small body size. We analyzed the differences in adult body size of males and females of the grasshopper Sphenarium histrio Gerstaecker (Orthoptera: Pyrgomorphidae) in three sites across an altitudinal gradient in southern Mexico. We also evaluated the possibility of protandry in these sampling sites using a common garden experiment. Male and female grasshoppers collected from low altitude sites in the field and reared in the laboratory were larger than those from a high altitude, suggesting genetic differentiation. Grasshoppers from a high altitude hatched earlier, had a shorter development time, presented fewer instars, and were smaller than grasshoppers from the other sampling sites. Moreover, development time in the three sampling sites was shorter in males than in females, suggesting protandry. Interestingly, the males from the three sites showed similar growth rates, but the females from low and high altitudes, respectively, had the fastest and slowest growth rates. In general, the adaptive value of the evolution of protandry has been focused on males. However, it may be that the growth rates of females in these sites could modify the degree of protandry as a response to their risk of pre‐reproductive death and the potential benefits associated with multiple matings.

Ecological notes on the Cát Bà gecko Goniurosaurus catbaensis, a species endemic to the limestone karst islands of northern Vietnam

The Cát Bà gecko Goniurosaurus catbaensis is endemic to Cát Bà and Ha Long islands of Vietnam. The ecology of this species is still scarcely studied but previous reports have suggested that the species is Endangered due to small population size and low ecological plasticity. We studied aspects of the ecology of this species between May 2019 and June 2020, along seven transects on Cát Bà island. At least 173 different individuals (possibly as many as 189) were recorded, with an equal sexratio and apparently no intersexual differences in adult body size. The species was clearly more widely distributed than previously supposed. We were able to confirm that G. catbaensis is a limestone karst microhabitat specialist, as the farthest sighting from a karst area was only 10 m. Geckos were observed at a mean height of 45 cm but up to 500 cm above ground, with males perching significantly higher than females. We observed these geckos at altitudes ranging from 11 to 228 m a.s.l.; much higher than previously recorded. Overall, our study revealed that this endemic gecko is certainly less threatened than previously feared.

Body size allometry impacts flight-related morphology and metabolic rates in the solitary bee Megachile rotundata

Body size is related to many aspects of life history, including foraging distance and pollination efficiency. In solitary bees, manipulating the amount of larval diet produces intraspecific differences in adult body size. The goal of this study was to determine how body size impacts metabolic rates, allometry, and flight-related morphometrics in the alfalfa leafcutting bee, Megachile rotundata. By restricting or providing excess food, we produced a range of body sizes, which allowed us to test the effect of body size on allometry, the power required for flight, and amount of energy produced, as measured indirectly through CO2 emission. The power required during flight was predicted using the flight biomechanical formulas for wing loading and excess power index. We found larger bees had higher absolute metabolic rates at rest and during flight, but smaller bees had higher mass-specific metabolic rates at rest. During flight, bees did not have size-related differences in mass-specific metabolic rate. As bees increase in size, their thorax and abdomens become disproportionately larger, while their wings (area, and length) become disproportionately smaller. Smaller bees had more power available during flight as demonstrated by flight biomechanical formulas. Smaller body size was advantageous because of a reduced power requirement for flight with no metabolic cost.

Captive breeding reveals larval performance and adult body size differences between two geographical populations of the stag beetle Aegus chelifer chelifer (Coleoptera: Lucanidae)

Stag beetles usually have great intraspecific variation in their body sizes, which can be affected by both environmental and genetic factors. However, direct studies on wild-caught specimens may be insufficient to clarify such variation due to the confounding effects of ecological variance in natural habitats. To evaluate this, the stag beetle Aegus chelifer chelifer MacLeay, 1819 was collected from within two localities (Bangkok metropolitan area and Chanthaburi province) in Thailand and then reared under the same condition to investigate the differences in morphological characteristics between the wild-caught and captive-bred beetles and between the two geographical populations. Narrow-sense heritabilities (h2) of the observed traits in adults were not significant. Variation in the body size of captive-bred specimens was less than in the wild-caught specimens and the overlap of the body size variation between the two populations was lower in the captive-bred beetles. The Chanthaburi population had a significantly larger body size than the Bangkok population. Allometric slopes and intercepts were also significantly different between the two geographic populations. Captive-bred larvae showed similar relative growth rates, but male larvae from the Chanthaburi population had a longer feeding period, and so a larger adult body size, than those from the Bangkok population. The differences between the two populations could be explained by adaptation through larval performances and body size in order to respond to their habitats.

Seasonal variations in body melanism and size of the wolf spider Pardosa astrigera (Araneae: Lycosidae).

Variations in species morphology and life‐history traits strongly correlate with geographic and climatic characteristics. Most studies on morphological variations in animals focus on ectotherms distributed on a large geographic scale across latitudinal and/or altitudinal gradient. However, the morphological variations of spiders living in the same habitats across different seasons have not been reported. In this study, we used the wolf spider, Pardosa astrigera, as a model to determine seasonal differences in adult body size, melanism, fecundity, and egg diameter both in the overwintering and the first generation for 2010 and 2016. The results showed that in 2010, both females and males of the overwintering generation were significantly darker than the first generation. Moreover, the overwintering females were markedly larger and produced more and bigger eggs than the first generation in both 2010 and 2016. Considering the overwintering P. astrigera experiencing low temperature and/or desiccation stress, these results suggest that substantially darker and larger body of the overwintering generation is adaptive to adverse conditions.

A new species of Salamander (Caudata, Plethodontidae, Bolitoglossa) from Serranía de los Yariguíes, Colombia

A new species of Bolitoglossa (Plethodontidae) is described from Andean cloud forests on the western slope of Serranía de los Yariguíes in the Cordillera Oriental of Colombia. This new species is distinguished from all other species of the Bolitoglossa adspersa species group by differences in adult body size, morphometric proportions, and tooth counts. The validity of the new species is also supported by molecular analyses. This new species increases the number of salamander species in Colombia to 23 and highlights the Cordillera Oriental of Colombia, with 11 species, as the region with the highest diversity of Bolitoglossa in South America.

Resource availability and sexual size dimorphism: differential effects of prey abundance on the growth rates of tropical snakes

Summary Broad phylogenetic patterns in sexual size dimorphism (SSD) are shaped by sex differences in net selection pressures (e.g. sexual selection, fecundity selection, survival selection), but environmental and ecological factors can also affect the expression of SSD. Discussions of proximate ecological influences on SSD have focused on niche divergence; for example, increase in a prey type used by only one sex can elevate growth rates of that sex but not the other. Food limitation also can generate spatial and temporal variation in SSD. Under restricted prey abundance, curtailed growth may mask SSD even if the optimal size is greater for one sex than the other. Because an increase in food availability elicits increased feeding and growth by the sex that benefits more from increased body size, variation in prey abundance can generate variation in SSD. We used mark‐recapture methods to study growth rates relative to prey (frog) abundance in two species of sexually dimorphic colubrid snake species in tropical Australia. In slatey‐grey snakes (Stegonotus cucullatus), a species in which larger body size enhances reproductive output in both sexes (because larger males win combat bouts, and larger females produce more/heavier eggs), increased abundance of frogs caused equivalent increases in growth rates in both sexes and hence did not affect SSD. In keelbacks (Tropidonophis mairii), a species in which larger size enhances reproductive output in females more than males (reflecting a lack of male–male combat), increased abundance of frogs elicited higher growth rates of females only. Thus, SSD in keelbacks was modified by prey abundance. Our results show that the magnitude of sex differences in adult body size can be influenced by proximate environmental factors and support the hypothesis of sex‐specific targets for maximum feeding rates. A lay summary is available for this article.

Sex hormones and the development of sexual size dimorphism: 5α-dihydrotestosterone inhibits growth in a female-larger lizard (Sceloporus undulatus).

Sexual differences in adult body size [sexual size dimorphism (SSD)] and color (sexual dichromatism) are widespread, and both male- and female-biased dimorphisms are observed even among closely related species. A growing body of evidence indicates testosterone can regulate growth, thus the development of SSD, and sexual dichromatism. However, the mechanism(s) underlying these effects are conjectural, including possible conversions of testosterone to estradiol (E2) or 5α-dihydrotestosterone (DHT). In the present study, we hypothesized that the effects of testosterone are physiological responses mediated by androgen receptors, and we tested two specific predictions: (1) that DHT would mimic the effects of testosterone by inhibiting growth and enhancing coloration, and (2) that removal of endogenous testosterone via surgical castration would stimulate growth. We also hypothesized that females share downstream regulatory networks with males and predicted that females and males would respond similarly to DHT. We conducted experiments on eastern fence lizards (Sceloporus undulatus), a female-larger species with striking sexual dichromatism. We implanted Silastic® tubules containing 150 µg DHT into intact females and intact and castrated males. We measured linear growth rates and quantified color for ventral and dorsal surfaces. We found that DHT decreased growth rate and enhanced male-typical coloration in both males and females. We also found that, given adequate time, castration alone is sufficient to stimulate growth rate in males. The results presented here suggest that: (1) the effects of testosterone on growth and coloration are mediated by androgen receptors without requiring aromatization of testosterone into E2, and (2) females possess the androgen-receptor-mediated regulatory networks required for initiating male-typical inhibition of growth and enhanced coloration in response to androgens.

Influence of in ovo mercury exposure, lake acidity, and other factors on common loon egg and chick quality in Wisconsin.

A field study was conducted in Wisconsin (USA) to characterize in ovo mercury (Hg) exposure in common loons (Gavia immer). Total Hg mass fractions ranged from 0.17 µg/g to 1.23 µg/g wet weight in eggs collected from nests on lakes representing a wide range of pH (5.0-8.1) and were modeled as a function of maternal loon Hg exposure and egg laying order. Blood total Hg mass fractions in a sample of loon chicks ranged from 0.84 µg/g to 3.86 µg/g wet weight at hatch. Factors other than mercury exposure that may have persistent consequences on development of chicks from eggs collected on low-pH lakes (i.e., egg selenium, calcium, and fatty acid mass fractions) do not seem to be contributing to reported differences in loon chick quality as a function of lake pH. However, it was observed that adult male loons holding territories on neutral-pH lakes were larger on average than those occupying territories on low-pH lakes. Differences in adult body size of common loons holding territories on neutral-versus low-pH lakes may have genetic implications for differences in lake-source-related quality (i.e., size) in chicks. The tendency for high in ovo Hg exposure and smaller adult male size to co-occur in low-pH lakes complicates the interpretation of the relative contributions of each to resulting chick quality.

Evolutionary divergence of adult body size and juvenile growth in sympatric subpopulations of a top predator in aquatic ecosystems.

Evolutionary theory predicts that different selective regimes may contribute to divergent evolution of body size and growth rate among populations, but most studies have focused on allopatric populations. Here, we studied five sympatric subpopulations of anadromous northern pike (Esox lucius) in the Baltic Sea subjected to allopatric habitats for a short period of their life cycle due to homing behavior. We report differences in adult body size among subpopulations that were in part due to variation in growth rate. Body size of emigrating juveniles also differed among subpopulations, and differences remained when individuals were reared in a common environment, thus indicating evolutionary divergence among subpopulations. Furthermore, a QST-FST comparison indicated that differences had evolved due to divergent selection rather than genetic drift, possibly in response to differences in selective mortality among spawning habitats during the allopatric life stage. Adult and juvenile size were negatively correlated across subpopulations, and reconstruction of growth trajectories of adult fishes suggested that body size differences developed gradually and became accentuated throughout the first years of life. These results represent rare evidence that sympatric subpopulations can evolve differences in key life-history traits despite being subjected to allopatric habitats during only a very short fraction of their life.

Altitudinally divergent adult phenotypes in Iberian wall lizards are not driven by egg differences or hatchling growth rates.

The interplay between ecological conditions and life histories has been widely acknowledged in vertebrates, particularly in lizards. Environmental conditions may exert different selective pressures and produce divergent phenotypes even in geographically and genetically close populations. The Iberian wall lizard constitutes a perfect model organism as it is considered a species complex with a complicated evolutionary history. Here, we focus on two proximate populations in which we examined adult morphology and reproductive investment of wild-caught lizards along a 500-m altitudinal gradient with contrasting environmental conditions, where adults show marked morphological differences in spite of being closely related. Also, we performed a common garden experiment to examine embryonic and hatchling growth. We focused on reproductive investment per clutch, incubation time, egg size, morphology and growth rate of hatchlings. Results showed clutch size differences between populations that were independent of the larger body size of highland females. However, there were no egg morphological differences between populations, except for egg width, and this difference disappeared after controlling for female body size. Hatchling lizards from both populations did not differ in morphology. Moreover, we did not observe differences between populations or sexes in hatchling growth. Overall, we provide evidence that the differences in adult body size and clutch size are not driven by size at hatching which is not contributed to by egg size, nor are intrinsic hatchling growth rates associated with the environmental conditions experienced in our common garden experiment, suggesting that adult phenotypes are not the result of intrinsic differences between populations.

Relationship between growth and standard metabolic rate: measurement artefacts and implications for habitat use and life-history adaptation in salmonids.

Mass-specific standard metabolic rate (SMR, or maintenance metabolism) varies greatly among individuals. Metabolism is particularly sensitive to variation in food consumption and growth creating the potential for significant bias in measured SMR for animals that are growing (e.g. juveniles) or of uncertain nutritional status. Consequently, interpreting individual variation in metabolism requires a sound understanding of the potentially confounding role of growth and the relative importance of fixed (genetic) vs. environmental drivers of SMR variation. We review the role of growth in measured SMR variation in juvenile salmonids, with the goals of (i) understanding the contribution of growth (and food consumption) to SMR variation through ontogeny, (ii) understanding the relative contributions of tissue maintenance and biosynthesis (overhead costs of growth) to apparent SMR variation, and (iii) using intrinsic growth effects on SMR to model how alternate life-history strategies may influence growth and measured SMR in juvenile salmonids. SMR measures on juveniles, even when post-absorptive, may be inflated by delayed growth-associated overhead costs, unless juveniles are on a maintenance ration (i.e. not growing). Empirical measurements of apparent SMR in food restricted vs. satiated 2-5g juvenile salmon demonstrate that estimates may be inflated by as much as 67% due to delayed overhead costs of growth, even when SMR measurements are taken 35h post-feeding. These results indicate that a substantial component of variation in apparent SMR among juvenile salmonids may be associated with (i) environmentally driven variation in ration (where elevated SMR measurements are an artefact of delayed growth overhead costs), (ii) intrinsic (genetic) or plastic organ-system trade-offs related to increasing investment in metabolically expensive digestive tissue responsible for processing food and (iii) intrinsic (genetic) variation in maximum body size and growth among individuals or life-history types. We suggest that selection for differences in adult body size among resident and anadromous forms leading to differences in juvenile growth trajectories may contribute to both SMR variation and habitat segregation in freshwater, where juveniles with higher growth are constrained to foraging in high velocity habitats to meet their greater consumption needs.

Mechanisms causing variation in sexual size dimorphism in three sympatric, congeneric lizards.

Sexual differences in adult body size (sexual size dimorphism, or SSD) ultimately can be favored by selection because larger males are more likely to be successful competitors for females, because larger females bear larger clutches, or because intersexual size differences reduce resource competition. Natural selection during juvenile development can influence sexual dimorphism of adults, and selection on adults and juveniles may differ. Studies that address the relative contributions of adult body shape dimorphism and sexually dimorphic patterns of growth and maturity are particularly useful in understanding the evolution of size dimorphism, yet they are rare. We investigated three sympatric, congeneric lizard species with different degrees and directions of adult sexual dimorphism and compared their growth patterns, survival probabilities, and intersexual trophic niche differences. Different mechanisms, even within these closely related, sympatric species, acted on juvenile lizards to produce species differences in adult SSD. Both degree and direction of dimorphism resulted from differences between the sexes in either the duration of growth or the rate of growth, but not from differences in rates of survival or selection on juvenile growth rate. Species- and sex-specific trade-offs in the allocation of energy to growth and reproduction, as well as differential timing of maturation, thus caused the growth patterns of the sexes to diverge, producing SSD. The differences that we observed in the direction of SSD among these species is consistent with their different social systems, suggesting that differential selection on adult body size has been responsible for the observed species-specific differences in juvenile growth rates and maturational timing.

Body-Size Evolution on Islands: Are Adult Size Variations in Tiger Snakes a Nonadaptive Consequence of Selection on Birth Size?

Mean adult size has been used as the traditional measure of body size to explain trends of insular gigantism and dwarfism in a wide array of taxa. However, patterns of variation in body size at birth have received surprisingly little attention, leaving open the possibility that adult body-size differences are nonadaptive consequences of selection acting on neonate body size. Here I used an empirical and correlative approach to test this hypothesis in a mosaic of 12 island and mainland snake populations in Australia. Data collected on 597 adult and 1,084 neonate tiger snakes showed that (1) both adult and neonate mean body sizes varied strongly across populations; (2) prey diversity and size convincingly explained birth-size variations: birth size-notably, gape size-correlated with prey size; (3) neonate snout-vent length was significantly correlated with neonate gape size; and (4) neonate snout-vent length was significantly correlated with adult snout-vent length. Postnatal growth rates recorded under common-garden conditions differed across populations and were correlated with mean prey size. These data collectively suggest that (1) prey size is the main driver for the evolution of body size at birth in gape-limited predators, (2) adult size variations may reflect selective forces acting on earlier life stages, and (3) adult size variations may also reflect resource availability during ontogeny (notably, prey diversity).

Host plant effects on morphometric characteristics of Liriomyza huidobrensis, L. sativae and L. trifolii (Diptera: Agromyzidae)

AbstractBody size of herbivorous insects may be directly related to host plant characteristics (e.g. nutrition, chemical composition) and positively linked to performance of the herbivore and its parasitoids. Differences in adult body size of Liriomyza huidobrensis, L. sativae and L. trifolii reared on Pisum sativum, Phaseolus vulgaris, Vicia faba and Solanum lycopersicum were assessed to determine the effect of host plants on adult size. Furthermore, morphometric wing and hind tibia measurements were used to determine suitability as proxy measures for adult size. Based on canonical and linear discriminant analyses, two distinct Liriomyza morphospecies groups were recognized across all plant species. The first cluster represented L. huidobrensis and the second L. sativae and L. trifolii. The overall degree of correct classification of the three species across all plants tested (correct observation) with cross validation was estimated at 85%. Liriomyza huidobrensis, the largest species, showed no differences in the variables measured when reared on the four plant species. However, different plant species seem to influence wing and hind tibia measurements in L. trifolii to a greater extent than in L. huidobrensis and L. sativae. The first two canonical variates could not separate the populations of L. huidobrensis and L. sativae reared on different plant species, but showed separation of L. trifolii populations into two clusters: firstly, insects reared on V. faba and secondly, insects reared on P. vulgaris and S. lycopersicum. Hind tibia length strongly correlated with wing length. Both can be used to determine adult size of the three Liriomyza species. This study provides some new evidence for the existence of differential preferences by Liriomyza species for the four host plant species. This, therefore, warrants larger scale field studies to test for positive preference‐performance of Liriomyza species on these plant species and to determine possible consequences at the third tritrophic level.

Proximate Contributions to Adult Body Size in Two Species of Dusky Salamanders (Plethodontidae: Desmognathus)

I used skeletochronological data to evaluate the contributions of propagule size, larval/juvenile growth, and age at first reproduction to differences in adult body size in two species of plethodontid salamanders of the genus Desmognathus. The traits in question were evaluated in populations of the larger D. quadramaculatus and smaller D. monticola in the southern Blue Ridge Mountains of North Carolina, USA. Gompertz and von Bertalanffy functions were fitted to the plots of standard length on skeletochronological age of each complete sample (larvae and/or juveniles, and adults) of earlier data sets; linear functions were fitted to data of immatures (larvae and/or juveniles). In order to examine the relationship of body mass and age, I conducted regressions of body mass on standard length in later samples of both species, calculated estimated body masses of the salamanders in the skeletochronological data sets, and then fitted a modified Gompertz function to each plot of body mass on age. The results showed that age at first reproduction is the principal factor contributing to differences between the species in size at first reproduction and adult size. Larger propagule sizes (i.e., hatching sizes) in D. quadramaculatus versus D. monticola may also be a factor; however, there is no evidence that a difference in larval/juvenile growth rate contributes to the adult size differential. Comparison of two populations of D. monticola suggested that small differences in adult body size result mainly from slight differences in age at first reproduction. Tradeoffs among propagule size, clutch size, growth, and age and size at first reproduction are discussed in the context of selective pressures that may have generated diversification in body size and life history in the genus.

Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from the Costa Rica-Panamá border region

Two new species of lungless salamanders (Plethodontidae) are described from high montane habitats of the border region between Costa Rica and Panamá. Bolitoglossa pygmaea and B. robinsoni are distinguished from each other and from other salamander species in this remote area by differences in adult body size, external proportions, foot webbing, tooth counts and/or external coloration. Both new species are assigned to the B. subpalmata species group, subgenus Eladinea. The miniaturized B. pygmaea is remarkable in being extensively depigmented, yet having the peritoneum and stomach area heavily pigmented and visible through the body wall.

Autecological effects of habitat alteration: trophic changes in mangrove marsh fish as a consequence of marsh impoundment

The key to predicting, counteracting, and managing environmental change is under- standing the mechanisms through which habitat change affects populations of organisms. The man- grove ecosystem of the Indian River Lagoon (IRL), east-central Florida, has been impounded for the purpose of mosquito control and is representative of such large-scale alterations. Sailfin mollies Poe- cilia latipinna in this system exhibit significant interpopulation differences in adult body size in rela- tion to habitat alteration. One likely hypothesis for this pattern is the apparent change in diet experi- enced as a result of impoundment, as has been documented in earlier studies. To address the question of whether these dietary changes would have been sufficient to cause trophic differences in P. latipinna populations relative to impoundment, a comparative study of trophic position in 2 size- divergent populations of P. latipinna using stable isotopes was employed. Results suggested that P. latipinna from an unimpounded site had a slightly higher trophic position than those from an impounded site and may constitute a partial explanation of differences in adult body size between the 2 populations. Results from the present study also showed that there was a fundamental differ- ence in energy source between the study sites, which suggests greater assimilation of mangrove- based carbon in impoundments. Specific population- and community-level effects of impoundment on mangrove habitats have been demonstrated, indicating a mechanistic link between anthro- pogenic habitat alteration and life history of P. latipinna.

Non‐lethal effects of invertebrate predators on Daphnia: morphological and life‐history consequences of water mite kairomone

Summary1. Here, we report morphological and life‐historical changes in the cladoceran Daphnia ambigua in response to chemical cues released by the predatory water mite Piona chilensis. Both species are common inhabitants of southern temperate lakes.2. We found significant differences in adult body size at first, second and third reproduction. Also, individuals exposed to kairomones had longer tail spines at first reproduction, and the resultant offspring had smaller bodies and shorter tail spines.3. Exposure to mite cues did not exert effects on brood size at first reproduction, but decreased offspring number in subsequent broods. Similarly, only the second and third reproduction events were delayed by kairomone exposure.4. The intrinsic population growth rate of predator‐induced animals was lower than that in controls, but simulations based on a parameterized matrix model showed that the fitness costs could be overcome if the reported phenotypic responses reduced predation rate moderately. The gain in protection from predators needed to cancel out the reduction in fitness associated with predator cues was directly related to juvenile survival and fertility, and inversely related to adult survival.5. This is the first work reporting phenotypic plasticity in Cladocera in response to kairomones released by water mites, which are conspicuous predators in many austral fresh waters.

The molecular forms of Anopheles gambiae: A phenotypic perspective

The African malaria mosquito Anopheles gambiae is undergoing speciation, being split into the M and S molecular forms. Speciation is the main process promoting biological diversity, thus, new vector species might complicate disease transmission. Genetic differentiation between the molecular forms has been extensively studied, but phenotypic differences between them, the evolutionary forces that generated divergence, and the mechanisms that maintain their genetic isolation have only recently been addressed. Here, we review recent studies suggesting that selection mediated by larval predation and competition promoted divergence between temporary and permanent freshwater habitats. These differences explain the sharp discontinuity in distribution of the molecular forms between rice fields and surrounding savanna, but they can also explain the concurrent cline between humid and arid environments due to the dependence on permanent habitats in the latter. Although less pronounced, differences in adult body size, reproductive output, and longevity also suggest that the molecular forms have adapted to distinct niches. Reproductive isolation between the molecular forms is achieved by spatial swarm segregation, although within-swarm mate recognition appears to play a role in certain locations. The implications of these results to disease transmission and control are discussed and many of the gaps in our understanding are highlighted.