Two methods of assessing the mortality factors affecting the larvae and pupae of Cameraria ohridella in the leaves of Aesculus hippocastanum in Switzerland and Bulgaria

The horse-chestnut leaf miner, Cameraria ohridella, is an invasive alien species defoliating horse-chestnut, a popular ornamental tree in Europe. This paper presents quantitative data on mortality factors affecting larvae and pupae of the leaf miner in Switzerland and Bulgaria, both in urban and forest environments. Two sampling methods were used and compared: a cohort method, consisting of the surveying of pre-selected mines throughout their development, and a grab sampling method, consisting of single sets of leaves collected and dissected at regular intervals. The total mortality per generation varied between 14 and 99%. Mortality was caused by a variety of factors, including parasitism, host feeding, predation by birds and arthropods, plant defence reaction, leaf senescence, intra-specific competition and inter-specific competition with a fungal disease. Significant interactions were found between mortality factors and sampling methods, countries, environments and generation. No mortality factor was dominant throughout the sites, generations and methods tested. Plant defence reactions constituted the main mortality factor for the first two larval stages, whereas predation by birds and arthropods and parasitism were more important in older larvae and pupae. Mortality caused by leaf senescence was often the dominant mortality factor in the last annual generation. The cohort method detected higher mortality rates than the grab sampling method. In particular, mortality by plant defence reaction and leaf senescence were better assessed using the cohort method, which is, therefore, recommended for life table studies on leaf miners.     

Life history of the sand fly vector Lutzomyia cruciata in laboratory conditions

Lutzomyia cruciata Coquillet (Diptera: Psychodidae: Phlebotominae) is a potential vector of Leishmania sp.; its geographical distribution in Mexico is widespread, but its life history is unknown. The present study gives relevant information on the life cycle, morphology, survival and reproduction of Lu. cruciata observed over successive generations under laboratory conditions. Seven successive generations were produced. A total of 975 adults were obtained in a sexual proportion of 1.1 : 1 (female : male). Each Lu. cruciata female produced 20.7 eggs and 1.9 adults, approximately, with a proportion of eggs per female of 2.7% (first generation) and 21.3% (second generation). The life cycle of Lu. cruciata, from egg to adult, occurred in 52.7 ± 0.52 days. The largest percentage of mortality occurred during the egg stage (48.5%) and the first larval instar (26.5%), whereas in the pupal stage mortality was the lowest (9.1%). Lutzomyia cruciata exhibits sexual dimorphism based on size, which is exhibited as of the second larval instar, males being smaller than females. The maximum survival of females and males was 10 and 15 days, respectively. An overview of the immature stages of the species made with an electronic scanning microscope is included. This paper contributes basic information on aspects of Lu. cruciata that were previously unknown related to its life history.     

Temperature thresholds and degree-day model for Marmara gulosa (Lepidoptera: Gracillariidae)

The developmental thresholds for Marmara gulosa Guillén & Davis (Lepidoptera: Gracillariidae) were investigated in the laboratory by using 17, 21, 25, 29, and 33 degrees C. The lowest mortality occurred in cohorts exposed to 25 and 29 degrees C. Other temperatures caused >10% mortality primarily in egg and first and second instar sap-feeding larvae. Linear regression analysis approximated the lower developmental threshold at 12.2 degrees C. High mortality and slow developmental rate at 33 degrees C indicate the upper developmental threshold is near this temperature. The degree-day (DD) model indicated that a generation requires an accumulation of 322 DD for development from egg to adult emergence. Average daily temperatures in the San Joaquin Valley could produce up to seven generations of M. gulosa per year. Field studies documented two, five, and three overlapping generations of M. gulosa in walnuts (Juglans regia L.; Juglandaceae), pummelos (Citrus maxima (Burm.) Merr.; Rutaceae), and oranges (Citrus sinensis (L.) Osbeck; Rutaceae), for a total of seven observed peelminer generations. Degree-day units between generations averaged 375 DD for larvae infesting walnut twigs; however, availability of green wood probably affected timing of infestations. Degree-day units between larval generations averaged 322 for pummelos and 309 for oranges, confirming the laboratory estimation. First infestation of citrus occurred in June in pummelo fruit and August in orange fruit when fruit neared 60 mm in diameter. Fruit size and degree-day units could be used as management tools to more precisely time insecticide treatments to target the egg stage and prevent rind damage to citrus. Degree-day units also could be used to more precisely time natural enemy releases to target larval instars that are preferred for oviposition.     

Adult diet affects the life history and host-killing behavior of a host-feeding parasitoid

Neochrysocharis formosa (Westwood), an important biocontrol agent of agromyzid leafminers worldwide, is a host-feeding, idiobiont parasitoid. Female wasps have three types of host-killing behaviors: reproductive (parasitism), non-reproductive host feeding (host feeding), and host stinging without oviposition or feeding (host stinging). In this study, we compared the life history and host-killing behaviors of female parasitoids under four adult diets: starvation, hosts only, hosts plus honey (10% w/v honey solution), and honey only. Furthermore, we analyzed the host-feeding and oviposition preferences of adult females in the hosts-only and hosts-plus-honey treatments. Female parasitoids feeding on hosts had significantly increased longevity, higher fecundity, more host-stinging events, and caused a higher total host mortality than parasitoids in the starvation treatment. The honey supplement significantly increased longevity, fecundity, host-stinging events, and total host mortality, as well as average daily fecundity, but did not alter host-feeding events, host-stinging events, or daily total host mortality. However, the honey supplement did reduce the number of daily host-feeding events and induced a shift toward oviposition. Finally, we found that the non-reproductive host killing caused by host feeding and host stinging enhanced the control potential of N. formosa. These results should contribute to a better understanding of the biocontrol efficiency of destructive host feeders.     

The biology and phenology of the citrus whitefly, Dialeurodes citri, on citrus in the Coastal Plain of Israel

The citrus whitefly, Dialeurodes citri (Ashmead) (Homoptera: Aleyrodidae) (CWF), is a polyphagous citrus pest, originating in S. E. Asia, which was first discovered in the Western Galilee region of Israel in 1975. The larval instars of CWF secrete honeydew, and the sooty mould fungus, which develops on the honeydew, causes damage to the tree and its fruit. In observations made in Israel between 1987 and 1991 CWF usually completed three generations per year, although a small proportion of the population had two or four generations. From October to April a combination of shortened photoperiod and reduced temperature arrest the development of CWF in the fourth larval instar. The average length of the life cycle from egg to adult emergence was 65 days in the first generation (April–June), 52 days in the second generation (July–August) and 8 months in the third generation (September–April). Under laboratory conditions of L16: D8 and 25 °C, the mean developmental time was 54 days, and the duration of the egg, first larval instar, second larval instar, third larval instar, fourth larval instar and putative pupa (red eyes) were 12.1, 6.5, 5.5, 7.6, 18, and 4.7 days, respectively. Adults prefer to lay their eggs on young, fully developed citrus leaves, but also oviposit on older leaves. The inner region and shaded sides of the tree are preferred for oviposition.     

Life Cycle of Culicospora magna (Kudo, 1920) (Microsporida: Culicosporidae) in Culex restuans Theobald with Special Reference to Sexuality1

The life cycle of Culicospora magna (Kudo, 1920) Weiser, 1977, consists of two major developmental sequences that alternate in host individuals of successive generations, each of the sequences starting with a sporoplasm and ending with spores. The first sequence occurs in larval, pupal, and adult stages of a parental generation of the host mosquito, Culex restuans Theobald; it begins with a sporoplasm from an ingested uninucleate spore and progresses through stages in gametogony, plasmogamy, nuclear association, merogony, karyogamy, and disporous sporulation with production of binucleate spores that discharge sporoplasms into the oocytes. The second sequence occurs in egg and larval stages of a filial generation of the same host species; it begins with the binucleate sporoplasm that entered the egg, includes stages in merogony, nuclear dissociation, and mictosporous sporulation, and ends with uninucleate spores. These spores are released into the environment following death of the host and are capable of infecting new parental generation host individuals. The life cycle is conceived as an alternation of generations related to haploidy and diploidy in the nuclei, the transition from haploidy to diploidy occurring with nuclear association and the transition from diploidy to haploidy occurring with nuclear dissociation.     

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16′ S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.     

两种共存网蛱蝶幼期的生命表研究

在河北省赤城县大海坨国家自然保护区内,从2002年到2004年应用生命表方法研究了在同一网络斑块中共存的金堇蛱蝶Euphydryas aurinia和大网蛱蝶Melitaea phoebe两个世代的幼期各阶段的死亡情况,目的是了解影响种群动态的重要因子,为它们的长期保育提供信息。结果表明,金堇蛱蝶幼期总累积死亡率都较小,两个世代分别为59%和72%; 而大网蛱蝶总累积死亡率较高,两个世代分别为89%和80%。影响大网蛱蝶死亡的最重要因子是放牧,两个世代与放牧相关的k值分别为0.559和0.167;尤其是在越冬后大网蛱蝶幼虫组聚集取食阶段,导致两个世代分别有50%和25%的幼虫组死亡。寄生蜂在大网蛱蝶小种群下也分别使两个世代4%和9%的5～6龄期幼虫以及13%和24%的蛹死亡。金堇蛱蝶死亡主要发生在越冬期,与越冬死亡相关的k值两个世代分别为0.073和0.199, 而寄主植物的质量影响越冬期幼虫组死亡; 寄生蜂则对金堇蛱蝶种群的调控作用极小,只有在2003～2004世代有4.0% 的越冬后幼虫被寄生和7%的蛹被寄生。影响两种蛱蝶种群动态的关键因子不同,采取的保护措施应有所不同。在春季减少源斑块内的放牧,对于以源-汇集合种群形式存在的大网蛱蝶种群恢复和增长十分必要; 而对以经典集合种群形式存在的金堇蛱蝶, 通过适当植被管理提高繁殖区域内寄主植物质量,可以提高越冬期幼虫组存活率,有利于其长期持续生存。     

Biology ofHyphantria cunea Drury (Lepidoptera: arctiidae) in Japan. V. Preliminary life tables and mortality data in urban areas

Studies on the population dynamics of the fall webworm, Hyphantria cunea have been carried out at three survey stations and along selected roads in the urban area of Tokyo since 1966. Twelve survivorship curves obtained during two years and 8 life tables show that the mortality rate in early developmental stages of the fall webworm is remarkably low as compared with that of other lepidopterous defoliaters and the mortality rate in later developmental stages is compensatory high. The low mortality rate in early stages is considered to be due to the protective role of the nest-web and the lack of egg and larval parasites. All but one parasitic species emerge from prepupae and pupae. Spiders living in the nest-web of the fall webworm play an important role in reducing the number of young larvae. Direct observations and caging experiments showed that relatively high mortality during later larval stages is mainly due to predation by birds (in the first generation) and wasps (in the second generation). The generation mortality in the survey stations always exceeded the level where the population is kept at the steady state, and the outbreak of this moth is considered to be continued by the immigration of adults from large trees growing in gardens on which the larvae can escape from predation pressure.     

Starvation induced stress and the susceptibility of the Colorado potato beetle,Leptinotarsa decemlineata,to infection by Beauveria bassiana

Starvation of second instar Colorado potato beetle larvae for 24h immediately after treatment with Beauveria bassiana conidia increased susceptibility to the pathogen and subsequent sporulation of cadavers but decreased time to larval death. In feeding studies, B. bassiana-treatment had no effect on subsequent larval development, and mortality occurred 5-6 days after treatment. Twenty-four hours of starvation alone retarded subsequent larval development but did not affect mortality. Mortality of B. bassiana-treated starvation stressed larvae occurred 4-5 days after treatment. Both B. bassiana treatment and 24h starvation significantly reduced total foliage consumption and daily weight gains. On the day of treatment, B. bassiana had no effect on the efficiency with which food was converted to biomass (ECI). ECI was not affected by B. bassiana or starvation alone on the day following treatment but was significantly affected by a combination of both. When larvae were exposed to a range of limited food quantities, ECI decreased with decreasing food availability but only extreme stress (starvation for 24h) increased susceptibility to B. bassiana. Topical application of Dacryodes excelsa resin (an antifeedant) to potato leaves caused a concentration dependent reduction in foliage consumption and weight gain by second instar larvae but did not affect larval mortality. When larvae were exposed to a fixed concentration of B. bassiana and a range of antifeedant concentrations there were significant linear relationships between 24h larval weight gain and mortality and 24h larval weight gain and sporulation. The interaction between starvation stress and the susceptibility to B. bassiana infection is discussed and its possible implications in pest management considered.     

Evaluation of lectin-expressing transgenic sugarcane against stalkborers (Lepidoptera: Pyralidae): effects on life history parameters

The impact of snowdrop lectin (Galanthus nivalis agglutinin, GNA) expressed in transgenic sugarcane on life history parameters of Mexican rice borer [Eoreuma loftini (Dyar)] and sugarcane borer [Diatraea saccharalis (F.)] (both Lepidoptera: Pyralidae) was evaluated. In the laboratory, lyophilized sugarcane leaf sheath tissue was incorporated in a meridic diet resulting in a GNA concentration of 0.47% of total protein, and used for insect bioassays over two successive generations. Deleterious effects of GNA were not observed on survival, weight, and developmental periods of larvae and pupae, nor on adult fecundity and egg viability of D. saccharalis. Moreover, in the first generation, addition of transgenic sugarcane tissue to the diet enhanced larval growth in D. saccharalis resulting in higher larval and pupal weight compared with diet with nontransgenic sugarcane, but this effect was not observed in the second generation. In contrast, larval survival, percent adult emergence, and female fecundity of E. loftini were significantly reduced when fed transgenic sugarcane diet compared with nontransgenic sugarcane diet. In addition, a substantial reduction of female pupal weight of E. loftini was observed in the second generation. For both species, the only consistent effect of GNA in both generations was a reduction in adult female longevity. Life table parameters showed that GNA at the level found in the transgenic diet negatively affected development and reproduction of E. loftini, whereas it had a nil to positive effect on development and reproduction of D. saccharalis.     

Population fluctuations of the diamondback moth,Plutella xylostella (L.) on cabbages inBacillus thuringiensis sprayed and non sprayed plots and factors affecting within-generation survival of immatures

Summary The population fluctuations and within-generation survival of immatures stages of the diamondback moth,Plutella xylostella on summer cabbages, were examined in unsprayed plots in 1984 and 1985 and in plots sprayed with two formulations ofBacillus thuringiensis in 1985. There was two distincy generations per crop and no noticeable difference in population fluctuations were observed in unsprayed plots between years. There was distinct difference between unsprayed and sprayed plots in 1985, from the larval period onwards. However, the survivorship patterns in all plots in both years were a Type 2 based onDeevey's (1947) classification. Life table studies showed that there was essentially no difference in the mortality agents acting on each of the stages except for numerical differences in the within-generation mortality rates. The major mortality rates during the egg stage were the parasitoid,Trichogramma spp. and unknown factors including rainfall; in the larval 1 stage was unknown mortality; in the larval 2 stage was the parasitoid,Apanteles plutellae and during the pupal stage was unknown mortality and parasitoid,Diadromus subtilicornis. Adult mortality was determined for generation 2. It was relatively higher in the unsprayed plots compared to theB. thuringiensis sprayed plots. The contribution of abiotic factors such as rainfall and temperature, and biotic factors such as parasitoids and predators in determining within-generation population levels and the fluctuation of populations on cabbage were discussed.     

Effects of GNA transgenic sugarcane on life history parameters of Parallorhogas pyralophagus (Marsh) (Hymenoptera: Braconidae), a parasitoid of Mexican rice borer

Parallorhogas pyralophagus (Marsh) is the principal parasitoid of the Mexican rice borer [Eoreuma loftini (Dyar)], the primary pest of sugarcane in south Texas. Insect resistant transgenic sugarcane producing Galanthus nivalis agglutinin (GNA) was developed to improve control of E. loftini. The present laboratory study addressed whether over two consecutive generations transgenic sugarcane delivered via artificial diet has adverse host-mediated effects on P. pyralophagus life history parameters. The results of this study showed that a number of life history parameters were affected by transgenic sugarcane, whereas others were not affected, and that effects varied between generations. In the first generation, adult longevity was increased by approximately 2 d, and cocoon to adult and egg to adult developmental times were prolonged by approximately 1 d in parasitoids exposed to transgenic sugarcane, whereas effects were not evident on adult size; egg load; egg to cocoon developmental time; rates of gain of longevity and egg load with adult size; and egg, larval, and pupal mortality. However, in the second generation, adult longevity was reduced by approximately 3 d, adult size by approximately 5%, egg load by approximately 24%, and rate of gain of longevity with adult size by approximately 21%, whereas effects were not evident on the rate of gain of egg load with adult size. It was concluded that although GNA transgenic sugarcane, ingested via E. loftini tissues, was not acutely toxic to P. pyralophagus, the sublethal effects on life history parameters measured in this study must be considered in a broader context to determine their possible ecological significance.     

Offspring performance in dynamic habitats: key factors for a riparian carabid beetle

Abstract.  1. Knowledge of the ecology of carabid species is largely restricted to adults, although larval mortality is assumed to be the key factor in overall mortality. As low-mobility larvae are unable to avoid unfavourable conditions, habitat selection of reproducing adults should be clearly affected by the habitat factors which determine offspring performance.
2. The present study examines the key habitat factors governing the distribution patterns of Bembidion velox larvae and adults on the river banks of the River Elbe by means of habitat suitability models. The validity of the determined habitat factors for offspring performance and survival was tested in laboratory experiments.
3. In the field, B. velox adults as well as larvae show a strong association with semi-terrestrial, sandy, open soil habitats.
4. In the laboratory, overall mortality of larvae reared in different substrates was lowest in sand of medium grain size mixed with fine and coarse sand. The first larval instars in particular reacted sensitively to variations in grain size. Furthermore, flood resistance of eggs was demonstrated, as 90% of larvae hatched under permanently flooded conditions.
5. Short development times were recorded, with 4–7 days for hatching of young larvae from eggs after oviposition and approximately 28 days for the development of adults from newly hatched larvae. This probably increases the survival probability for the pre-imaginal stages in dynamic habitats.
6. In conclusion, it can be said that reproductive success is strongly dependent on oviposition site selection by adults as this reflects the ecological demands of the immature stages.     

Laboratory evaluation of Alsystin, a chitin synthesis inhibitor, against Culex pipiens pipiens L. (Dip., Culicidae): effects on development and cuticle secretion

Alsystin, a benzoylphenylurea derivative, was evaluated on Culex pipiens pipiens L. Treatment was made on newly third- and fourth-instar larvae for 24 h. Mortality occurred at various developmental stages following treatment. However, death as larvae was relatively important compared with the mortality recorded for the other later developmental stages. Treatment resulted in a dose-dependent reduction in adult emergence. Moreover, treatment caused morphological aberrations depending upon the importance of larval–adult transformation and significantly increased the duration of both third and fourth larval instars. A histological study conducted on fourth instar larval integument, revealed that Alsystin delayed the ecdysis and significantly reduced the thickness of both larval and pupal cuticles secreted compared with controls. Moreover, the apolysis that occurred at day 4 in controls was not affected by treatment.     

Mortality of the terrestrial adult and aquatic nymphal life stages of Baetis vernus and Baetis rhodani in the Breitenbach, Germany (Insecta: Ephemeroptera)

1. The mortality of Baetis vernus Curtis and Baetis rhodani Pictet during the terrestrial-aerial and aquatic life stages was studied at the Breitenbach near Schlitz, Hesse, Germany. The number of females emerging from the stream was recorded with emergence traps. To estimate mortality of females of both species during terrestrial life, numbers of emerging females were compared with numbers of females returning to the stream for oviposition, as shown by numbers of egg masses found in the stream. 2. Mortality of female B. vernus during their terrestrial life stage was 98.8%. It was 91.2 and 96.6%, respectively, during the first and second generations of B. rhodani. 3. To estimate the mortality of both sexes of B. rhodani during the aquatic life stage, the number of eggs laid by the first generation was compared with the number of adults emerging during the second generation. 4. Mortality during the aquatic phase (both sexes combined) of B. rhodani was 91.1%. 5. Mortality during the terrestrial life stages was at least as high, if not higher than during the aquatic stages. Evidently, there is a considerable export of organic material into the terrestrial environment around the stream. Mortality during terrestrial life may be an important regulator of population size.     

Variability in the life-history patterns of four species of Hydropsyche (Trichoptera: Hydropsychidae) in southern Ontario streams

The life-history patterns of four Hydropsyche species (Trichoptera: Hydropsychidae) found in the Credit and Humber Rivers of southern Ontario were investigated through rearing studies, larval and adult collections, and analyses of densities and mortality of field populations of pupae. Hydropsyche slossonae was univoltine but was probably capable of producing a second generation under favourable conditions at the warmest station at which it occurred. Hydropsyche sparna was partially bivoltine at upstream stations and bivoltine at downstream stations. At downstream stations, the common species were Hydropsyche bronta . which was trivoltine. and Hydropsyche morosa , which was bivoltine. Hydropsyche sparna, H. bronta and H. morosa all exhibited a split-cohort development, in which not all of the summer-produced larvae completed development, pupated and emerged before the end of the growing season. Overwintering populations consisted of young larvae that had hatched from eggs laid by adults of the previous generation. Analysis of the temporal changes in the densities of living and dead pupae provided a reliable way to separate overwintering and spring- and summer-produced generations.     

Shifting preference between oviposition vs. host-feeding under changing host densities in two aphelinid parasitoids

Destructive host-feeding is common in hymenopteran parasitoids. Such feeding may be restricted to host stages not preferred for oviposition. However, whether this is a fixed strategy or can vary according to resource levels or parasitoid needs is less clear. We tested the trade-off between host feeding and oviposition on two whitefly parasitoids under varying host densities. Females of two aphelinid parasitoids, Eretmocerus hayati and Encarsia sophia were exposed to nine different densities of their whitefly host, Bemisia tabaci, in single-instar tests to identify their functional response. Mixed-instar host choice tests were also conducted by exposing whiteflies at four densities to the parasitoids. We hypothesized that the parasitoid females can detect different host densities, and decide on oviposition vs. host-feeding accordingly. The results showed that both Er. hayati and En. sophia females tended to increase both oviposition and host-feeding with increased host density within a certain range. Oviposition reached a plateau at lower host density than host-feeding in Er. hayati, while En. sophia reached its oviposition plateau at higher densities. At low densities, Er. hayati parasitized most on first and second (the optimal ones), and fed most on third nymphal instars (the suboptimal one) of the whitefly host as theory predicts, while at high densities, both parasitism and host-feeding occurred on first and second instars which are preferred for oviposition. En. sophia parasitized most on third and fourth (the optimal ones), while fed on first instars (the suboptimal one) at low densities, and utilized third and fourth instars for both at high densities. In conclusion, oviposition vs. host-feeding strategy of parasitoid females was found to vary at different host densities. The balance between reserving optimal hosts for oviposition or using them for host-feeding depended on parasitoid life history and the availability of host resources.     

Artificial selection on larval growth curves in Tribolium: correlated responses and constraints

Body size is often constrained from evolving. Although artificial selection on body size in insects frequently results in a sizable response, these responses usually bear fitness costs. Further, these experiments tend to select only on size at one landmark age, rather than selecting for patterns of growth over the whole larval life stage. To address whether constraints may be caused by larval growth patterns rather than final size, we implemented a function‐valued (FV) trait method of selection, in which entire larval growth curves from Tribolium were artificially selected. The selection gradient function used was previously predicted to give the maximal response and was implemented using a novel selection index in the FV framework. Results indicated a significant response after one generation of selection, but no response in subsequent generations. Correlated responses included increased mortality, increased critical weight, and decreased development time (DT). The lack of response in size and development time after the first generation was likely caused by increased mortality suffered in selected lines; we demonstrated that the selection criterion caused both increased body size and increased mortality. We conclude that artificial selection on continuous traits using FV methods is very efficient and that the constraint of body size evolution is likely caused by a suite of trade‐offs with other traits.