Sensitivity to juvenile hormone is not reduced in clustering monarch butterflies, Danaus plexippus, in Australia

ABSTRACT. Female monarch butterflies, Danaus plexippus (L.) were collected from clusters in the Sydney region at weekly intervals after cluster formation. They were neck-ligated and injected with juvenile hormone (JH) in order to test for any period of lowered JH sensitivity during the non-reproductive clustering phase. Response was assessed by production of mature oocytes. In all weekly samples, the mean response was equal to or greater than that of newly emerged females reared and maintained in optimal conditions, and was substantially higher than that reported for diapausing monarchs in North American clusters. The result is further evidence for the absence of a true reproductive diapause in New South Wales populations.     

Energy reserves and metabolic expenditures of monarch butterflies overwintering in southern California

Abstract. 1. Energetic expenditure and predicted requirements for overwintering metabolism were determined for monarch butterflies ( Danaus plexippus L.) in southern California.
2. Fat content of butterflies declined steadily from a maximum of 71% lean dry weight in late November to a minimum of 36% lean dry weight in late January. The energy expenditure measured by fat depletion over a 61 day period from 24 November to 25 January was 26.05 joules per day.
3. Butterflies were generally the same temperature as the air at any time they were inactive, whether they were part of a large cluster or roosting solitarily.
4. Oxygen consumption of butterflies increased in a curvilinear fashion with increasing air temperature. Thus, the predicted metabolic requirements for an inactive butterfly during their quiescent period from late November to late January was 30.32 joules per day.
5. In contrast to the steady depletion of fat reserves during their quiescent period in December and January, low and stable fat levels of butterflies in late February coincide with high levels of flight activity, mating and emigration of females from the colony.
6. Progressive tightening of the energy balance due to increasing nocturnal temperatures from northern to southern California coastal areas may limit the southern distribution and duration of overwintering aggregations.     

Do Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected Monarch Butterflies Overwintering in Mexico

Long-distance migration can lower parasite prevalence if strenuous journeys remove infected animals from wild populations. We examined wild monarch butterflies (Danaus plexippus) to investigate the potential costs of the protozoan Ophryocystis elektroscirrha on migratory success. We collected monarchs from two wintering sites in central Mexico to compare infection status with hydrogen isotope (δ ²H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ ²H values than parasitized butterflies, indicating that uninfected butterflies originated from more northerly latitudes and travelled farther distances to reach Mexico. Within the infected class, monarchs with higher quantitative spore loads originated from more southerly latitudes, indicating that heavily infected monarchs originating from farther north are less likely to reach Mexico. We ruled out the alternative explanation that lower latitudes give rise to more infected monarchs prior to the onset of migration using citizen science data to examine regional differences in parasite prevalence during the summer breeding season. We also found a positive association between monarch wing area and estimated distance flown. Collectively, these results emphasize that seasonal migrations can help lower infection levels in wild animal populations. Our findings, combined with recent declines in the numbers of migratory monarchs wintering in Mexico and observations of sedentary (winter breeding) monarch populations in the southern U.S., suggest that shifts from migratory to sedentary behavior will likely lead to greater infection prevalence for North American monarchs.     

Biological Observations of Monarch Butterfly Behavior at a Migratory Stopover Site: Results from a Long-term Tagging Study in Coastal South Carolina

Like most migratory species, monarch butterflies (Danaus plexippus) must stop frequently during their long southward migration to rest and refuel, and the places where they stop are important for the success of the migration. The behavior of monarch butterflies at migratory stopover sites has never been examined in detail. Here we present results of a long-term study of monarchs at one stopover site in coastal South Carolina where over 12,000 monarchs have been captured, measured and tagged (with numbered stickers to track recovery rates) over 13 years. Only 3 monarchs (0.023%) were recovered at the monarchs’ overwintering sites in Mexico, which is consistent with other tagging studies on the eastern coast. The migration season was longer at this site than at inland locations and monarchs continued to be captured in November and December, when most monarchs had already arrived at the overwintering areas in Mexico. In addition, there were 94 monarchs captured between Jan 1 and Mar 15, indicating that some monarchs overwinter at this site. Of all monarchs captured during the migration season, 80% were captured while nectaring and 10% while roosting. Others were basking, resting, flying and even mating. The sex ratio was male biased by three to one in all behavior categories except those captured mating. Roosting and nectaring monarchs had fresher wings than those in other behavior categories, suggesting that these are younger individuals. There were 13 observations of females ovipositing on non-native Asclepias curassavica during the fall months, which speaks to the potential for this plant to pull monarchs out of the migratory pool. Aside from these insights, this study also serves as an example of the potential that monarch tagging studies have to advance scientific understanding of monarch migration.     

Oviposition by Danaus plexippus in relation to cardenolide content of three Asclepias species in the southeastern U.S.A.

Abstract. 1. Female monarchs were observed in the field ovipositing on a native North American milkweek host, Asclepias humistrata L. As in a comparable Australian study on an introduced novel host ( Asclepias fruticosa L.) we found post-alighting rejection of plants with low and high cardiac glycoside concentration (CG).
2. Most oviposition took place on plants with CG in the range 200–500 μg/0.1 g dry weight. Thin-layer chromatography showed no obvious qualitative difference in cardenolide types between accepted and rejected plants, excepting an indication that rejected plants may have a higher level of more polar cardenolides.
3. In a controlled laboratory experiment comparing oviposition on a low ( A.incarnata L.) vs high ( A.curassavica L.) CG host plant species we found no relationship between CG and oviposition on the low CG species, but a negative relationship in the high CG species. This corroborates our findings on oviposition on single host species in the field.
4. We also record the first indication of a physiological cost of monarch larvae feeding on plants high in CG. There was a significant negative relationship between survival of first instar larvae and CG levels in plants. This study opens the way for further work on the association of monarch butterflies and their toxic milkweed hosts.     

Local and cross‐seasonal associations of climate and land use with abundance of monarch butterflies Danaus plexippus

Quantifying how climate and land use factors drive population dynamics at regional scales is complex because it depends on the extent of spatial and temporal synchrony among local populations, and the integration of population processes throughout a species’ annual cycle. We modeled weekly, site‐specific summer abundance (1994–2013) of monarch butterflies Danaus plexippus at sites across Illinois, USA to assess relative associations of monarch abundance with climate and land use variables during the winter, spring, and summer stages of their annual cycle. We developed negative binomial regression models to estimate monarch abundance during recruitment in Illinois as a function of local climate, site‐specific crop cover, and county‐level herbicide (glyphosate) application. We also incorporated cross‐seasonal covariates, including annual abundance of wintering monarchs in Mexico and climate conditions during spring migration and breeding in Texas, USA. We provide the first empirical evidence of a negative association between county‐level glyphosate application and local abundance of adult monarchs, particularly in areas of concentrated agriculture. However, this association was only evident during the initial years of the adoption of herbicide‐resistant crops (1994–2003). We also found that wetter and, to a lesser degree, cooler springs in Texas were associated with higher summer abundances in Illinois, as were relatively cool local summer temperatures in Illinois. Site‐specific abundance of monarchs averaged approximately one fewer per site from 2004–2013 than during the previous decade, suggesting a recent decline in local abundance of monarch butterflies on their summer breeding grounds in Illinois. Our results demonstrate that seasonal climate and land use are associated with trends in adult monarch abundance, and our approach highlights the value of considering fine‐resolution temporal fluctuations in population‐level responses to environmental conditions when inferring the dynamics of migratory species.     

Lack of genetic differentiation between monarch butterflies with divergent migration destinations

Monarch butterflies are best known for their spectacular annual migration from eastern North America to Mexico. Monarchs also occur in the North American states west of the Rocky Mountains, from where they fly shorter distances to the California Coast. Whether eastern and western North American monarchs form one genetic population or are genetically differentiated remains hotly debated, and resolution of this debate is essential to understand monarch migration patterns and to protect this iconic insect species. We studied the genetic structure of North American migratory monarch populations, as well as nonmigratory populations in Hawaii and New Zealand. Our results show that eastern and western migratory monarchs form one admixed population and that monarchs from Hawaii and New Zealand have genetically diverged from North American butterflies. These findings suggest that eastern and western monarch butterflies maintain their divergent migrations despite genetic mixing. The finding that eastern and western monarchs form one genetic population also suggests that the conservation of overwintering sites in Mexico is crucial for the protection of monarchs in both eastern and western North America.     

Effects of photoperiod, temperature, and host plant age on induction of reproductive diapause and development time in Danaus plexippus

Abstract 1. Diapause induction in monarch butterflies Danaus plexippus was studied using adults captured from the wild in Minnesota and Wisconsin, and individuals reared under outdoor and controlled conditions.
2. Oocyte presence in females and ejaculatory duct mass in males were used to indicate reproductive status.
3. Some wild individuals were in diapause in mid-August, and all males and females were in diapause by late August and early September respectively.
4. Individuals reared under decreasing day lengths and fluctuating temperatures were more likely to be in diapause than were individuals reared under long or short day lengths or constant temperatures.
5. Individuals fed potted old Asclepias curassavica plants were more likely to be in diapause than were those fed potted young host plants; when cuttings of Asclepias syriaca plants from the field or greenhouse pots were used, there was no effect of host plant age.
6. Extremely high temperatures increased the number of day-degrees required for development from egg to adult, while decreasing day lengths and older host plants tended to decrease the number of day-degrees required for development.
7. There appears to be a continuum of reproductive development in monarchs, with gradual declines in mean ejaculatory duct mass and oocyte production during the late summer.
8. None of the experimental treatments led to 100% diapause, and diapause was more likely to occur in monarchs subjected to more than one diapause-inducing cue.     

Ecological niches in sequential generations of eastern North American monarch butterflies (Lepidoptera: Danaidae): the ecology of migration and likely climate change implications

Eastern North American monarch butterflies (Danaus plexippus L.) show a series of range shifts during their breeding season. Using ecological niche modeling, we studied the environmental context of these shifts by identifying the ecological conditions that monarchs use in successive summer months. Monarchs use a consistent ecological regimen through the summer, but these conditions contrast strikingly with those used during the winter. Hence, monarchs exhibit niche-following among sequential breeding generations but niche-switching between the breeding and overwintering stages of their annual cycle. We projected their breeding ecological niche onto monthly future climate scenarios, which indicated northward shifts, particularly at the northern extreme of their summer movements, over the next 50 yrs; if both monarchs and their milkweed host plants cannot track these changing climates, monarchs could lose distributional area during critical breeding months.     

Monarch butterflies reared under autumn-like conditions have more efficient flight and lower post-flight metabolism

1. Many migratory animals undergo physiological and behavioural changes to prepare for and sustain long-distance movements. Because insect migrations are common and diverse, studies that examine how migratory insects meet the energetic demands of long-distance movements are badly needed. 2. Monarch butterflies (Danaus plexippus) migrate up to 4000 km annually from eastern North America to wintering sites in central Mexico. Autumn generation monarchs undergo physiological and behavioural changes in response to environmental cues to initiate migration. In particular, exposure to cooler temperatures and shorter day lengths in early autumn causes monarchs to enter the hormonally induced state of reproductive diapause. 3. This study examined differences in flight-associated metabolic rate (MR) and flight performance metrics for monarchs experimentally reared under autumn-like conditions (typically experienced before the southward migration) relative to monarchs reared under summer-like conditions. 4. Adult monarchs reared under autumn-like conditions showed lower post-flight MRs, greater flight efficiency, and lower measures of reproductive activity relative to monarchs reared under summer-like conditions. Increases in post-flight metabolism were associated with monarch body weight, age, and flight velocity. 5. These findings suggest that a trans-generational shift in flight energetics is an important component of the monarch's complex migratory syndrome, and that physiological changes that accompany reproductive diapause facilitate energy conservation during flight.     

Host plant pattern and variation in climate predict the location of natal grounds for migratory monarch butterflies in western North America

The breeding grounds of migrant generation monarch butterflies in eastern North America are well known. In stark contrast the location of natal grounds of western migrants has not been delineated. We show that 55% of the area within seven western states was potential breeding range based on: (1) the occurrence of milkweed host plant species with phenology making them available during late-summer and (2) regional thermal conditions supportive of adult reproductive activity and development of immature stages. We next used a series of spatially explicit “bottom-up” regression models to test this first-approximation natal origins distribution. We tested for associations between variation in moisture availability at putative natal habitat and inter-annual variation in monarch abundance at western wintering sites for a 10 year period (1998–2007). Variation in moisture availability, as measured by Palmer’s drought severity index (PDSI), across the western region predicted monarch abundance patterns. In contrast and as expected, PDSI across known eastern breeding grounds did not predict variation in western monarch migrant abundance. The pattern of moisture availability was not uniform between states or within states and permitted similar tests of association at a finer geographical level. PDSI for California, Idaho, Nevada, and Oregon (but not Arizona, Utah, or Washington) were each significantly associated with monarch wintering abundance patterns with California exhibiting the strongest relationship. At a more focused spatial scale we tested the local recruitment hypothesis. This is the notion that western coastal wintering monarch populations derive only from nearby coastal breeding habitat and that monarchs do not migrate from more distant natal grounds. Variation in moisture availability within a block of three contiguous central California climate divisions (Sacramento Drainage, San Joaquin Drainage, and Southeast Desert Basin) significantly predicted inter-annual abundance of migrant generation monarchs. In contrast PDSI patterns of three coastal California climate divisions, i.e., ones local to wintering sites, as well as that of climate divisions in western Nevada and Arizona did not predict variation in monarch abundance at this more focused spatial resolution. Our findings suggest that moisture regimes act as a strong bottom-up driver of monarch abundance pattern via resource availability in western USA.     

Behavioral and ecological interactions of foraging mice (Peromyscus melanotis) with overwintering monarch butterflies (Danaus plexippus) in México

Summary Mice (Peromyscus melanotis) immigrate extensively to overwintering colonies of monarch butterflies (Danaus plexippus) in México. There they feed on both live and dead butterflies that accumulate on the ground and in low vegetation. Through a series of feeding experiments, we examined the potential impact of mouse predation on these colonies, as well as how this predation was influenced by the accessibility and the degree of desiccation of the monarchs. Mice attacked on average 39.9 wet (freshlykilled) butterflies per night. We estimated that a population of mice (75–105 individuals) could kill approximately 0.40–0.57 million butterflies in a 1 ha colony (4–5.7% of the colony) over the 135-day overwintering season. In feeding experiments, mice fed disproportionately on: 1) wet (hydrated) monarchs close to the ground versus those perched higher; 2) wet monarchs, when both wet and dry (desiccated) monarchs were on the ground; and 3) wet monarchs on stakes versus dry monarchs on the ground. Mice commonly ate the entire abdomen of dry monarchs, whereas they fed selectively on the abdomen of wet monarchs by discarding the bitter, cardenolide-laden cuticle and eating the internal tissues. These results suggest that the monarchs' state of desiccation is more important than their accessibility in determining the feeding preferences of these mice. However, the monarchs' strong tendency to crawl up vegetation does appear to reduce their risk to mouse predation.     

Stable isotopes (δD and δ13C) are geographic indicators of natal origins of monarch butterflies in eastern North America

Wing membranes of laboratory and field-reared monarch butterflies (Danaus plexippus) were analyzed for their stable-hydrogen (δD) and carbon (δ¹³C) isotope ratios to determine whether this technique could be used to identify their natal origins. We hypothesized that the hydrogen isotopic composition of monarch butterfly wing keratin would reflect the hydrogen isotope patterns of rainfall in areas of natal origin where wings were formed. Monarchs were reared in the laboratory on milkweed plants (Asclepias sp.) grown with water of known deuterium content, and, with the assistance of volunteers, on native milkweeds throughout eastern North America. The results show that the stable hydrogen isotopic composition of monarch butterflies is highly correlated with the isotopic composition of the milkweed host plants, which in turn corresponds closely with the long-term geographic patterns of deuterium in rainfall. Stable-carbon isotope values in milkweed host plants were similarly correlated with those values in monarch butterflies and showed a general pattern of enrichment along a southwest to northeast gradient bisecting the Great Lakes. These findings indicate that natal origins of migratory and wintering monarchs in Mexico can be inferred from the combined δD and δ¹³C isotopic signatures in their wings. This relationship establishes that analysis of hydrogen and carbon isotopes can be used to answer questions concerning the biology of migratory monarch butterflies and provides a new approach to tracking similar migratory movements of other organisms. Received: 1 July 1998 / Accepted: 11 November 1998     

Freeze-protection of overwintering monarch butterflies in Mexico: critical role of the forest as a blanket and an umbrella

Abstract. 1. At their high-altitude overwintering sites in Mexico, monarch butterflies frequently are subjected to sub-zero°C temperatures during December-March. Although monarchs have moderate supercooling ability, two ecological factors strongly influence their capacity to resist freezing: wetting and exposure to the clear night sky. 2. As shown in Fig. 2, 50% of a population of butterflies with water on their body surfaces freeze at warmer sub-zero temperatures (-4.2°C) compared to butterflies with no water on their bodies (-7.7°C). 100% mortality occurs, respectively, at ?7.7°C and ?15°C. 3. Comparative measurements of rainfall within a large overwintering colony in Mexico indicated that the intact canopy acts as an umbrella that reduces butterfly wetting during winter storms. 4. Variable experimental exposure of butterflies to the clear night sky indicated that openings in the forest canopy increases radiational cooling and causes monarch body temperatures to drop as much as 4°C below ambient air temperature. Monarchs under dense cover had body temperatures approximately the same as the ambient air temperature, but more exposed individuals had body temperatures below ambient in direct proportion to the degree of exposure. Consequently, forest thinning increases the probability that the butterflies will freeze to death. 5. Whereas both wetting and exposure are increased by disturbance of the forest canopy, the interaction of these two factors exacerbates freezing mortality during winter storms: 50% of dry and unexposed butterflies froze at ?8°C, whereas wetted and fully exposed butterflies froze at only ?0.5°C. 6. Butterflies inside and on the bottom of the fir bough clusters are better protected from wetting than those on the outside. This supports the hypothesis that the structure of the butterfly clusters has evolved through individual selection to avoid wetting. 7. The data strongly reinforce previous evidence that forest thinning should be totally prevented within and adjacent to the overwintering sites in order to minimize both wetting and exposure of the butterflies that synergistically increase winter mortality at the overwintering sites in Mexico.     

REPRODUCTIVE DIAPAUSE IN TYPHLODROMUS OCCIDENTALIS NESBITT (ACARINA: PHYTOSEIIDAE) FROM SOUTHERN NEW SOUTH WALES

Abstract
Studies on the reproductive diapause of an introduced insecticide-resistant strain of the phytoseiid predator, Typhlodromus occidentalis Nesbitt, from southern New South Wales are reported. Diapause was induced in 43–59% of predators when reared at 16 or 19°C under photophases of 10–12 h. No diapause induction response curve with changing photophase was evident. Diapause in overwintering predators was readily terminated by exposure to warm temperatures and a long photophase. The rapidity of diapause termination increased progressively during winter. The apparent loss of dependency upon reproductive diapause as an overwintering strategy in this strain of T. occidentalis is discussed.     

Embryonic mortality and predation on egg capsules of the Port Jackson shark Heterodontus portusjacksoni (Meyer)

Egg capsules of the Port Jackson shark Heterodontus portusjacksoni were examined during underwater visual census surveys at two sites in New South Wales, Australia, during the 2002–2005 reproductive seasons (austral winter). Embryonic mortality was high (0·783–0·896 per annum) with the majority (99·2%) resulting from predation. The crested horn shark Heterodontus galeatus and common tent shell Astralium tentoriformis (Gastropoda, Turbinidae) were positively identified as egg predators. Gastropod predation was insignificant, accounting for only 2·0 and 2·1% of total mortality at Terrigal Haven and Dent Rock, respectively. Most capsules appeared to have been depredated by large predatory fishes with the eastern blue groper Achoerodus viridis (Labridae) and the black stingray Dasyatis thetidis (Dasyatidae) as possible candidate predators. The rate of embryonic mortality in H. portusjacksoni is higher than that reported for other oviparous elasmobranchs. This high level of mortality has significant consequences for the conservation and management of this species, especially when combined with an understanding of their low fecundity, late maturity and protracted reproductive life.     

Monarchs in Australia: On the Winds of a Storm?

The monarch butterfly, Danaus plexippus, is one of Australia's best-known exotic butterflies, being first recorded here in the spring/summer of 1870/1871. However, the source of the original population is unknown. Using historical records we suggest that the most likely source of the founder population was from Vanuatu and/or New Caledonia. Many almost simultaneous 'first records' for the butterfly in Australia suggest that a large, well-distributed population was present when first noticed. While such a population may have developed from a limited number of individuals flying across the Coral Sea, the well documented, very dramatic appearance of large monarch populations in Australia does not appear to fit this model. Rather, we hypothesise that large numbers of monarchs were carried to Australia on cyclonic winds: no fewer that 3 cyclones hit the Queensland coast in early 1870. If one or more of these cyclones tracked from the Vanuatu/New Caledonia chain, then they may have transported monarchs. Once established on the central/northern Queensland coast, natural migration would account for the appearance of butterflies further south. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cardiac glycosides and oviposition by Danaus plexippus on Asclepias fruticosa in south-east Queensland (Australia), with notes on the effect of plant nitrogen content

Abstract. 1. Previous studies which attempt to relate oviposition by female monarchs, Danaus plexippus (L.), to the cardiac glycosides of their milkweed host plants are reviewed and found to be defective.
2. Here we followed ovipositing females in the field and recorded their oviposition behaviour on individual Asclepias fruticosa (L.) plants. The cardenolide glycoside and per cent nitrogen of each of these plants and also of a control set of plants was recorded.
3. Milkweeds accepted by females, on which egg laying took place, had significantly lower mean cardenolide levels compared with rejected plants and a control sets of plants.
4. Most oviposition ( c. 70%) took place on plants with intermediate cardenolide (CG) levels (=200–300 μg/0.1 g DW).
5. The mean CG level in A.fruticosa was 345 μg/0.1 g DW (SD = 138, n =122, range 119–719).
6. There was no relationship between monarch oviposition behaviour and plant per cent nitrogen.     

Description and Biology of A New Species of Cybocephalus Erichson (Coleoptera: Nitidulidae), A Predator of Australian Citrus Whitefly

Cybocephalus aleyrodiphagus sp.n., a predator of Australian citrus whitefly, Orchamoplatus citri (Takahashi), is described from adults and larvae collected from southern New South Wales (where it appears to be the major natural enemy of O. citri ) and South Australia. Eggs of C. aleyrodiphagus are laid singly near egg circles of O. citri and larvae feed on eggs, crawlers and nymphs. Adults feed on all immature stages of O. citri. Reproductive, but not non-reproductive, females contain greater quantities of stearic, oleic and linoleic acid than males indicating the possible presence of a sex pheromone. In southern New South Wales, C. aleyrodiphagus sp.n. overwinters as non-reproductive adults and produces two adult generations during December-January and March-May. the sex ratio is close to 1:1 during October-December (overwintered adults) but male biased (2.2:1) during January-July (new generation adults). Trends of abundance for C. aleyrodiphagus correspond well with those for O. citri nymphs/pupae indicating a functional response of the predator to prey, despite disruption by foraging ants.     

Reward Tracking and Memory Decay in the Monarch Butterfly,Danaus plexippus L. (Lepidoptera: Nymphalidae)

Due to their long‐distance migration routes and high longevity, monarch butterflies (Danaus plexippus) are likely to benefit from learning how to discriminate and remember suitable feeding resources. In this study, we assessed monarchs’ abilities to track changing nectar sources over time and to retain learned information presented in two conditioning schedules. Non‐preferred (blue and red) and preferred (yellow) artificial flowers were concomitantly offered to monarchs in a three‐phase experiment. In each phase, flowers of only one color contained sucrose solution, while the others contained water. The rewarding color was changed in each phase. Instantaneous observations were made to assess butterfly visits to each color during each phase; continuous observations over the first 90 min of a new phase allowed us to look in more detail at the transition process. Overall, monarchs tracked sucrose availability, visiting the rewarding flowers more often than the unrewarding ones, regardless of innate preferences. However, butterflies reverted to innate color preferences when the newly rewarding color was different from the initial trained color. In a second experiment, memory decay was compared for butterflies trained according to two schedules: ‘single training’ (sucrose solution in red vs. water in blue artificial flowers in one 15‐min session per day) or ‘intermittent training’ (as above, but in two 7.5‐min sessions per day). Afterwards, butterflies were tested on alternate days for a week in arrays containing unrewarding models of both colors. Following either training schedule, memory persisted for at least 3 d after reinforcement ceased. Our findings reveal that monarchs are able to change their feeding responses according to the flowers’ reward status despite innate preferences, as well as to retain flower information for about half a week regardless of the conditioning dynamics.