Heat Production in the Voodoo Lily (Sauromatum guttatum) as Monitored by Infrared Thermography

The pattern of surface temperatures of the inflorescence of Sauromatum guttatum was investigated by using an infrared camera. The male flowers are weakly thermogenic on the first day of inflorescence opening (D-day) as well as on the next day (D + 1), reaching 0.5 to 1°C above ambient temperature. The appendix (the upper sterile part of the inflorescence) is highly thermogenic on D-day, reaching 32°C, and is faintly thermogenic on D + 1, reaching 1°C above ambient temperature. The lower part of the spadix, close to the female flowers, is also thermogenic on D-day and D + 1, reaching a temperature similar to that of the appendix only on D + 1. Salicylic acid does not induce heat production in the lower part of the spadix, as it does in the appendix. Respiration of tissue slices obtained from the appendix shows that the capacity for cyanide-insensitive respiration is present in young and mature appendices. This alternative respiratory pathway is not, however, utilized in young appendix tissue, but is engaged during the maturation of that tissue.     

HEAT-PRODUCTION AND CROSS-POLLINATION OF THE ASIAN SKUNK CABBAGE SYMPLOCARPUS RENIFOLIUS (ARACEAE)

The Asian skunk cabbage Symplocarpus renifolius has an exothermic spadix on which about 100 flowers bloom in very early spring when effective pollinators such as bees and drone flies are inactive. This species is protogynous; female phase and male phase took 6.8 ± SD 5.8 days and 16.7 ± 5.7 days, respectively, with a short transitional phase of bisexuality (2.1 ± 0.9 days). The spadices produced heat 24 hours/day throughout female and bisexual phases, but temperature dropped quickly after the beginning of male phase. Although self-compatibility was expected from the flower structure, the basipetal flowering, and the absence of effective pollinators, bagging tests demonstrated that they rarely produce seeds without crossing. The spadices were visited by small numbers of invertebrates throughout the flowering season. Of these invertebrates, house flies, rove beetles, and mosquitos were the likeliest pollinators, since they are probably attracted both to the pollen produced in male phase and to the stench or carbon dioxide in female phase. On two female spadices with immature male flowers, we fortuitously collected a rove beetle and a mosquito that carried some pollen grains; these had to have been transported from other S. renifolius spadices. This infrequent and ineffective pollination appears to explain why as low as 13% of spadices set seeds in a natural population. We examine alternative hypotheses to explain production of heat in spadices of skunk cabbage.     

Thermogenesis and flowering biology of <Emphasis Type="Italic">Colocasia gigantea</Emphasis>, Araceae

The thermogenesis and flowering biology of Colocasia gigantea (Blume) Hook. f. were studied from December 2005 to February 2006 on Espiritu Santo, Vanuatu (South Pacific). Endogenous thermogenesis was measured in two ways: (1) continuously over 5-day periods, and (2) over 3 h during maximum heating. The study showed that heat was generated by the male part of the spadix and probably the lower zone of the sterile region. The temperatures of the male part peaked twice: (1) between 0625 and 0640 (during the female phase) and (2) 24 h later (during the male phase). The average maximum temperature was 42.25 ± 0.14°C during the female phase (16.63°C above the ambient temperature) and 35.14 ± 0.22°C during the male phase (10.61°C above the ambient temperature). In the lower zone of the sterile region, thermogenesis was documented only during the female phase. The average maximum temperature was 35.44 ± 0.41°C (9.82°C above the ambient temperature). Thermogenic heating appeared to be closely associated with the activities of pollinating insects.     

Pollination mutualism between Alocasia macrorrhizos (Araceae) and two taxonomically undescribed Colocasiomyia species (Diptera: Drosophilidae) in Sabah, Borneo

Two taxonomically undescribed Colocasiomyia species were discovered from inflorescences of Alocasia macrorrhizos in Kota Kinabalu City, Sabah, Borneo, Malaysia. The aims of this study were to investigate the reproductive ecology of the flies and the plant, ascertain the importance of the flies as pollinators and examine the intimate association between flowering events and life history of the flies. We conducted sampling, observations and field pollination experiments. The flies were attracted by the odour of female‐phase inflorescences in the early morning on the first day of anthesis. They fed, mated and oviposited in the inflorescences for 1 day. On the second day, the flies, covered with pollen grains, left the male‐phase inflorescences for the next female‐phase inflorescences. The immature forms of both fly species hatched, developed and pupated within the infructescences without damaging the fruits, and developed adults emerged when the mature infructescences dehisced. The flowering events and fly behaviours were well synchronized. In field pollination experiments, inflorescences bagged with a fine mesh (insect exclusion) produced almost no fruits, whereas those bagged with a coarse mesh (bee exclusion) produced as many fruits as the open‐pollinated controls. These results indicate that these flies are the most efficient and specialised pollinators for their host, A. macrorrhizos. These flies, in return, depend on A. macrorrhizos for food and habitat through most of their life cycle. This study provides a deeper insight into the less recognised, highly intimate pollination mutualism between Araceae plants and Colocasiomyia flies.     

Homeosis, morphogenetic gradient and the determination of floral identity in the inflorescences ofPhilodendron solimoesense (Araceae)

A comparative developmental study of the inflorescence ofPhilodendron solimoesense was conducted using scanning electron microscopy. The spadix ofP. solimoesense is characterized by unisexual flowers. Staminate flowers are initiated on the upper portion of the spadix while pistillate flowers develop on the lower portion of the spadix. An intermediate zone located between the upper male and lower female portion of the inflorescence consists of sterile male flowers. Within this intermediate zone a row of flowers exhibit polarity with respect to the identity of sexual organs. Stamens are initiated on the flank of the floral meristem facing the upper male zone and carpels are initiated on the portion of the floral meristem facing the lower female zone. The resulting flowers therefore assume a bisexual identity. At the level of the inflorescence, all floral buds are initiated along a series of contact parastichies and the continuity of these parastichies is not disrupted at any level in the male, intermediate, and female zones on the spadix. Results from this study support the presence of a morphogenetic gradient acting at the level of the inflorescence and appears to be independent of the boundaries of floral primordia.     

The cowl does not make the monk: scarab beetle pollination of the Neotropical aroid Taccarum ulei (Araceae: Spathicarpeae)

Taccarum ulei (Araceae, Spathicarpeae) is a seasonal geophytic aroid, native to north‐eastern Brazil, that flowers during two months of the rainy season. Patterns of floral thermogenesis, pollination biology, and floral traits associated with pollination syndromes were studied and compared with those of other Araceae. Two species of cyclocephaline scarabs (Scarabaeidae, Cyclocephalini) were recognized as effective pollinators: Cyclocephala celata and Cyclocephala cearae. Larvae of an unidentified species of fruit fly (Melanoloma spp., Richardiidae, Diptera) were also frequently observed in inflorescences at various maturation stages, feeding on the connectives of male florets and fruits, and thus lowering the reproductive success of individual plants. Beetles were attracted by odoriferous inflorescences in the early evening of the first day of anthesis, during the female phase. The emission of attractive volatiles was coupled with intense thermogenic activity in the entire spadix, unlike other aroids in which only certain zones of the spadix heat up. Pollen release, which marks the beginning of the male phase on the subsequent evening, was not related to floral thermogenesis. Comparative multivariate analysis of the floral traits of T. ulei points to a beetle‐pollinated aroid, although some of the observed traits of the species are not common to other taxa sharing this pollination strategy. Such incongruence might be explained by the evolutionary history of the tribe Spathicarpeae and potential pollinator shifts. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Studying Lords and Ladies,Arum maculatum (L.)

Historical uses of Arum maculatum (L.) are summarized. Field and laboratory investigations show plants with purple and pale spadices are equally attractive to flies of the genus Psychoda; and these are attracted to the scent in the absence of visual stimuli. Over 75% of the flies which enter spikes escape. Temperature changes during spadix maturation, and spathe uncurling in the presence of flower tissue, indol-3-yl acetic acid, and gibberellic acid are investigated in the laboratory. Dilute solutions of both phytohormones cause significant spathe uncurling. Further investigations, suitable for project work in schools and colleges, are suggested.     

Function of the heater: the dead horse arum revisited

The dead horse arum, Helicodiceros muscivorus, is a conspicuous, foul smelling and thermogenic plant of the Araceae family. This Mediterranean arum lily copies several aspects of a carcass in order to attract carrion blowflies, which are subsequently exploited as unrewarded pollinators. We have previously shown that this plant exhibits a highly accurate olfactory carrion mimicry, which serves to attract the blowflies. In this study, we have investigated the role of thermogeny in the arum. We show that the thermogeny has a direct effect on the pollinators, altering their behaviour. By manipulating heat and odour release of the plant, we can show that the heat, produced along the appendix, is important to lure the flies to this structure, which is vital as the flies from the appendix are more prone to enter the trap chamber that houses the female and male florets. This study provides rare evidence for a direct functional role of thermogeny.     

Life history traits related to resource partitioning between synhospitalic species of Colocasiomyia (Diptera, Drosophilidae) breeding in inflorescences of Alocasia odora (Araceae)

Abstract.

• 1 Two closely related species of Colocasiomyia alocasiae (Okada) and C.xenalocasiae (Okada) (Diptera, Drosophilidae) breed in inflorescences of Alocasia odora C. Koch (Araceae), a hermaphroditic understorey clonal herb.
• 2 The two drosophilid species form a synhospitalic pair in Okinawa with alocasiae breeding in the upper half of the inflorescence and xenalocasiae breeding in the lower half.
• 3 C.alocasiae also has the following combination of life history traits: small body size, many eggs, and early reproductive maturity. In contrast, xenalocasiae can be described as having larger body size, fewer eggs, and delayed reproductive maturity.
• 4 Resource partitioning between the two species on the same host may be affected by these life history traits which are associated with their larval habitats.

     

Scent chemistry and pollinator attraction in the deceptive trap flowers of Ceropegia dolichophylla

Ceropegia species (Apocynaceae, Asclepiadoideae) have pitfall flowers and are pollinated by small flies through deception. It has been suggested that these flies are attracted by floral scent. However, the scent that is emitted from Ceropegia flowers has not been studied using headspace and gas chromatography mass spectrometry methods. It has also been unclear whether or not the flowers are mimics of particular models that attract flies. In the present study, we determined the composition as well as the spatial and temporal patterns of floral scent emitted by C. dolichophylla. Furthermore, we determined the pollinators in the native (China) and non-native (Germany) range of this species, and tested the capability of the floral scent to attract flies in the non-native range. Our data demonstrate that the floral scent, which is emitted from morning until evening, primarily from the tips of the corolla lobes, consists mainly of spiroacetals and aliphatic compounds. Milichiid flies were common visitors/pollinators in the native as well as non-native range, and were attracted by floral scent in bioassays performed in the non-native range. The compounds emitted by C. dolichophylla are unusual for flowers, but are well known from insect pheromones and occur in the glandular secretions of insects. The milichiid flies that visit and pollinate the flowers are kleptoparasites that feed on the prey (haemolymph or other secretions) of predatory arthropods, e.g. spiders, to which they are attracted by scent. Our data thus suggest that the floral scent of C. dolichophylla mimics the feeding sites of kleptoparasitic flies.     

Energy Loss in Tissue 2Sauromatum guttatum (Schott) Analysed by Microcalorimetry

The heat evolved by 1 mm-thick tissue slices of the appendixof the Sauromatum guttatum inflorescence was measured calorimetricallyduring development. From D–5 (5 d before inflorescence-opening,designated as D-day) to D–2 about 8 µW mg^–1fresh wt. was observed, and during D–1 an increase inheat evolution to 14 µW mg^–1 fresh wt. was monitored.The heat was produced through oxidative metabolism, since replacingthe air in the microcalorimeter with nitrogen blocked heatproduction.Addition of salicylic acid to tissue slices of thermogenic organs(appendix, lowest part of the spadix and male flowers) and ofnon-thermogenic organs (female flowers, club-shaped organs andlower part of the spadix) of Sauromatum inflorescences revealedthat the acid boosted heat-production only in the thermogenicorgans. The effect of the acid manifested itself with no appreciablelag time, and it generated non-linearity in the rate of heat-productionby tissue slices of the appendix. In the appendices of the highlythermogenic Arum italicum and of the weakly thermogenic Amorphophallusrivieri, salicylic acid selectively boosted the rate of heat-productionin the appendix of A. italicum. A Q₁₀ of 24 was found between15C to 25 C for 1 mm-thick slices of D–4 appendicesof S guttatum. Addition of digitonin or deoxycholate to pre-D-daytissue slices of the appendix increased the rate of heat-production. Key words: Amorphophallus rivieri, Arum italicum, microcalorimetry, salicylic acid, Sauromatum guttatum     

Different regulation of leaf respiration between Spinacia oleracea, a sun species, and Alocasia odora, a shade species

Mature leaves of shade species exhibit lower respiratory rates than those of sun species. To elucidate the mechanism underlying different respiratory rates between sun and shade species, we examined respiratory properties of leaves in Spinacia oleracea L., a sun species, and Alocasia odora (Lodd.) Spach, a shade species, with special reference to changes in the respiratory rate throughout the night. In S. oleracea , rates of both CO₂ efflux and O₂ uptake decreased with time during the night, whereas in A. odora both rates were virtually constant at lower levels. The rates of O₂ uptake in S . oleracea increased upon addition of sucrose, and the rates attained were virtually identical throughout the night. However, the addition of an uncoupler [carbonyl cyanide p -(trifluoromethoxy)-phenylhydrazone; FCCP] did not alter the rates. In contrast, the rates of O₂ uptake in A. odora were enhanced by the addition of FCCP, but not by sucrose. The concentrations of carbohydrates in the tissue decreased throughout the night in both species and the ATP/ADP ratio was always greater in A. odora. These results indicate that, in S. oleracea , the availability of respiratory substrate determines the respiratory rate, while the low respiratory rate in A. odora is ascribed to its low demand for ATP.     

Thermogenesis of three species of Arum from Crete

Inflorescences of arum lilies have a three-part spadix with a scent-producing, sterile appendix above two bands of fertile male and female florets. The appendix and male florets are thermogenic, but with different temporal patterns. Heat-production was measured in Arum concinnatum , A. creticum and A. idaeum . The male florets of A. concinnatum showed a 3 d continuous episode of thermogenesis with three waves, and the appendix warmed in a single, 6 h episode. Maximum fresh-mass-specific CO₂ production rate was 0.17  µ mol s⁻¹ g⁻¹ to achieve a 10.9 °C temperature elevation by the appendix, and 0.92  µ mol s⁻¹ g⁻¹ to achieve a 4.8 °C elevation by male florets. Reversible, physiological temperature regulation was not evident in either tissue. Respiration increased with tissue temperatures with Q₁₀ values of 1.8–3.9, rather than less than 1.0 as occurs in thermoregulatory flowers. Experimental step changes in temperature of appendix and male floret tissues also failed to show thermoregulatory responses. The patterns of thermogenesis therefore appear to be fixed by the temporal sequence of blooming. Thermogenesis in the alpine species, A. creticum and A. idaeum , was significantly lower than in the lowland A. concinnatum , possibly related to difficulty in raising floral temperature in their cold and windy habitat.     

Zonal thermogenetic dynamics of two species of Philodendron from two different subgenera (Araceae)

Spadix temperature was measured in two species of Philodendron : P. melinonii (subgenus Philodendron ) and P. solimoesense (subgenus Meconostigma ). For each species, the temperature of the male zone, the sterile male zone and the female zone of the spadix were recorded. In both species, the temperature of the male zone warmed up at the beginning of each of the two flowering nights. In P. melinonii , the temperature of the male sterile zone increased the first day but remained not significantly different from that of the ambient air during the second day. The temperature of the male zone warmed up slightly on the second day. In P. melinonii , the temperature of the three zones was not significantly different from that of the ambient air between the two peaks. In P. solimoesense , the temperature of the male zone and sterile zone rose to above that of the ambient air during the first night and then progressively cooled down but remained 3–6°C above the ambient air temperature until the second peak. In both species the temperature of the female zones remained more or less constant during the entire flowering cycle, very close to the temperature of the ambient air. We suggest that the heat production and the spadix temperature patterns observed may reflect a general physiological process common to all species of Philodendron . The biphasic pattern present in the subgenus Meconostigma can be seen as a variant of the 'two peaks' pattern, occurring in the subgenus Philodendron , with a 'plateau' phase between them. The comparison of the different thermogenic cycles occurring in Philodendron , Arum and Dracunculus seems to indicate some clear evolutionary trends. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 79–86.     

濒危植物南川升麻传粉生物学的研究

南川升麻（ＣｉｍｉｃｉｆｕｇａｎａｎｃｈｕａｎｅｎｓｉｓＨｓｉｓｏ）的繁育系统以异交为主,传粉昆虫主要为蝇类,盛花时的访花频率平均每花每天１１次,雄蕊脱落后约０６次;单穗升麻（Ｃ．ｓｉｍｐｌｅｘＷｏｒｍｓｋ）以虫媒为主,风也可以帮助其花粉在近距离传播。单穗升麻的传粉昆虫有蝇类和蜂类,盛花时的访花频率蝇类为１．４次,蜂类为１．２次。南川升麻和单穗升麻都是靠大的花序和气味来吸引传粉昆虫,报酬是退化雄蕊基部分泌的蜜液。自然条件下南川升麻每个柱头上平均有４枚花粉,结籽率平均为６４％,而单穗升麻每个柱头上平均有１７枚花粉,结籽率平均为４５９％。南川升麻有性生殖中传粉的限制是其致濒的重要因素之一。     

Pollination Biology in Cimicifuga nanchuanensis,an Endangered Species (Ranunculaceae)

The pollination biology of Cimicifuga nanchuanensis Hsiso, an endangered species and C. siraplex Wormsk, a widespread species as a control, has been studied. Protandry occurs in both species, the former goes further than the latter. The breeding system of C. nanchuanensis is dominantly xenogamous. The average visitation rate of flies as the main pollinators is 1.1 and 0.6 per flower per day at anthesis and after stamen shedding respectively. C. simplex is mainly an entomophilous plant. However, wind pollination could occur within a short distance. Moreover, a certain ratio of self pollination exists also in this species. The mean visitation rates of flies and bees, the main pollinators of C. simplex, are 1.4 and 1.2 respectively at anthesis. The pollinating insects of both plant species are attracted by their big inflorescences and the fragrant odor. Nectar, the reward for pollinators, produces in the secretory cells at the base of the spoonform staminode. In natural conditions, an average of 17 pollen grains are found on the surface of a stigma resulting in a mean seed set rate of 45.9 %. Pollination limitation (very few pollinators and their low efficiency of pollination) in C. nanchuanensis is one of the weak links in its life cycle and one of the reasons for the endangerment of this species.     

Pollination ecology of Arum italicum (Araceae)

The pollination ecology of Arum italicum was studied in south-western France. This plant attracts olfactory dung-breeding flies through deceit. These insects are principally represented by Diptera, all belonging to saprophyte families. The volatilization of the odouriferous compounds, responsible for their attraction, is achieved through the production of heat by the appendix. The insects are trapped for 24 h in order to participate in both sexual phases of the protogynous inflorescence. The male flowers produce three heat events during flowering. These peaks of heat seem to be involved in the spathe movements, since they occur during the opening of the inflorescence and the liberation of the insects. The last male heat event may be linked with the liberation of pollen and its dispersion by stimulating trapped flies. According to their frequency and pollen-load, two Psychoda species appear to be the most efficient pollinators ( P. crassipenis and P. pusilla ). Nevertheless, each of the other attracted species could play a significant role under different spatio-temporal conditions. Experiments on self-pollination have shown that obligate cross-pollination is necessary for A. italicum to set seeds. Moreover, hand- and natural-pollinated plants showed similarly high abortion frequencies suggesting that seed set may be more constrained by resources rather than by pollination limitation.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 205–214.     

Effects of GF-120 fruit fly bait concentrations on attraction, feeding, mortality, and control of Rhagoletis indifferens (Diptera: Tephritidae)

Effects of different concentrations of GF-120 NF Naturalyte Fruit Fly Bait on attraction and feeding responses, mortality, and control of the western cherry fruit fly, Rhagoletis indifferens Curran, were determined. In the laboratory, flies that had been exposed to sugar and yeast extract and then deprived of all food for 16-20 h were attracted to 40.0% GF-120, but not to 0.6 and 4.8% GF-120 (vol:vol). Nonstarved flies were not attracted to any concentration. Flies in the field were not attracted to 55.6% GF-120 on cherry leaves, and few flies fed on the bait. In the laboratory, males fed for shorter durations on and ingested lower amounts of 0.6% than 4.8 or 40.0% GF-120, but females fed equally on all concentrations. Spinosad in GF-120 was highly toxic to flies. Lethal concentrations50 (LC50 values) of spinosad for starved flies at 1-4 d were 1.5-0.7 ppm. When gravid flies were exposed to cherries treated with 0.6, 4.8, and 40.0% GF-120, mortality was greater at each higher concentration, but none prevented oviposition. Field spray tests comparing 0.6, 4.8, and 40.0% GF-120 in 225 ml of spray per cherry tree resulted in 79-94% lower larval infestations than in controls, but no differences were seen among the concentrations. Evidence from this study indicates that fresh 40.0% GF-120 was attractive in the laboratory but that flies were not attracted to fresh GF-120 from far distances within trees, suggesting that suppression of populations is caused in large part by flies finding the bait through normal movement over large areas.     

榕果挥发物对传粉榕小蜂的吸引作用

榕树 /榕小蜂专一性共生系统的维持 ,与榕树开花期释放的特殊的挥发性化合物以及榕小蜂对其寄主榕树的化学识别和定位紧密相关。研究选取了西双版纳地区常见的 3种榕树 ,即对叶榕 Ficus hispida、木瓜榕 F.auriculata和鸡嗉子榕F.semicordata的榕果作为实验材料 ,利用野外诱捕实验、室内生物检测实验检测传粉榕小蜂 Hymenoptera:ChalcidoidaeAgaonidae对 12种信息化合物及榕果的二氯甲烷浸提物的趋向性反应 ,研究不同榕属植物的传粉榕小蜂对相同的信息化合物的反应差异 ,以及传粉榕小蜂受不同发育时期榕果浸提物吸引的显著性程度。诱捕实验中对叶榕小蜂 Ceratosolen solmsimarchali对香叶醇的趋向性反应显著 ,大果榕小蜂 C.emarginatus对接受期榕果浸提物和芳樟醇都有明显的趋向性反应 ,而对间花期榕果浸提物则无显著反应。嗅觉仪生物检测实验中 ,鸡嗉果榕小蜂 C.gravelyi对香叶醇和松油醇都表现出显著的趋向性反应。结果表明 ,对叶榕、鸡嗉子榕传粉榕小蜂对 12种信息化合物的反应存在一定的差异 ,木瓜榕传粉榕小蜂对香叶醇和木瓜榕接受期榕果浸提物的趋向性反应比间花期榕果强得多     

Development of an odour-baited target for female New World screwworm, Cochliomyia hominivorax: studies with host baits and synthetic wound fluids

Field studies were conducted in Mexico to investigate the possibility of replacing sentinel animals as baits for female New World screwworm Cochliomyia hominivorax (Coquerel) (Diptera: Calliphoridae) with a synthetic odour-bait composed of compounds identified from wounds infested with the larvae of C. hominivorax. Studies using sheep held in stanchions surrounded by an incomplete ring of electrified nets showed that both male and female C. hominivorax were attracted to an unwounded animal (ratio of male : female flies caught 1.11 : 1.00), although they were not observed to land. By contrast, catches associated with unwounded sheep with larval wound fluid applied to shaven shoulders suggested that female C. hominivorax were attracted to (male : female ratio of electrified nets catch 0.14 : 1.00) and landed on the wound fluid but male flies did not respond (male : female ratio alighting on fluid 0.05 : 1.00). In related studies the attractiveness of synthetic blends of 25 electrophysiologically active compounds identified in fluids associated with wounds infested with the larvae of C. hominivorax were tested. The acidic components of the wound fluids were found to attract and elicit a landing response from both male and female flies, whereas the non-acidic components alone caught only low numbers of flies. However, the numbers of male and female flies that were attracted to and landed on a synthetic bait could be increased significantly by increasing the proportion of non-acid to acid components in the lure. In some replicates the most effective blend caught a number of C. hominivorax comparable to that caught by a standard synthetic attractant, Swormlure-4, although, unlike with Swormlure-4, the catch was predominately composed of female flies. It is uncertain whether Swormlure-4 contains compounds that elicit a landing response from male C. hominivorax or whether the synthetic and natural wound odours contain compounds that inhibit a landing response from male C. hominivorax. Further work is required to simplify the blend of compounds needed to attract female flies and to better understand the role of the compounds that elicit behavioural responses from both male and female C. hominivorax before a synthetic substitute can be fully developed to replace sentinel animals.