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MOSQUITO PHYTOPHAGY, POLLINATION AND
EVOLUTIONARY TRANSITION TO HAEMATOPHAGY
MADHUSUDAN M
PALB-8138
Sr. M.Sc. Agril. Entomology 1
PHYTOPHAGY:
Phytophagy - feeding on the plant or/and plant derived
products.
DO FEMALE MOSQUITOES FEED
ON PLANT/ PLANT DERIVED
PRODUCTS?
Yes
2
DIFFERENT TYPES OF RESOURCES OF PLANTS
 Floral nectar
 Extra-floral nectar
 Honey dew
 Plant tissue
 Ant regurgitates
 Rotting fruits
3
WHAT MAKES THEM(FEMALES) BLOOD FEEDERS?
4
ARE THERE ANY ADVANTAGES OF SUGAR
FEEDING?
 Sugar feeding is a fundamental characteristic of
mosquito life
 Energetically, sugar and blood are interchangeable
 The option to take sugar is retained for the competitive
advantage
 Even a replete blood meal does not initiate
development if reserves become too low as a result of
sugar deprivation
 Sugar feeding enhances longevity and
 Delayed egg laying (enhance fitness by being choosy)
5
6
Effect of sugar feeding on Anopheles gambiae on
survival and biting frequencies.
Water only Sugar + BloodBloodSugar solution
15151515
7
Mating
X 5
Gary and Foster, 2001
Females
only water
Males
10% sucrose
Males
10% sucrose
Females
only water
Mating
8
Sugar solution Blood Sugar + Blood
1 X 20 cage 1 X 20 cage1 X 20 cage
Gary and Foster, 2001
Mean survival times of female Anopheles
gambiae maintained on one of four diets
9
Age-specific survivorship (lx) of Anopheles gambiae
females on four diets when females were either (A) pooled
15 to a cage or (B) isolated individually. Gary and Foster, 2001
Proportion of Anopheles gambiae females
biting each day, with access either to blood
only or to blood + sugar
10
Gary and Foster, 2001
 Plant source include both
-carbohydrate (major)
-non carbohydrate
 Among all the sources-
Floral nectar is the major source
 But the content and nutrient status varies from source
to source, and plant species to plant species
11
Are plants provide only sugar (CHO) to mosquitoes?
12
(A)Impatiens walleriana,
(B) Asclepias curassavica,
(C) Passiflora edulis,
(D) Campsis radicans leaf,
(E) C. radicans green seed-pod,
(F) Ricinus communis.
Positive control
10% sucrose in a 1.5 ml
tube stuffed with cotton
Negative controls
1. a drip irrigated soil pot,
2. a cotton-stuffed tube of water
3. an empty cage
X 5
10 males +
10 females
Effect of different nectar sources on
survival of Aedes aegypti
Chen and Kearney, 2015
13
days
Proportion of mosquitoes surviving on different host
plants and controls
Chen and Kearney, 2015
FEEDING ON PLANT TISSUE-
IS IT A COMMON PHENOMENA?
(A) Anopheles sergentii probing on a leaf.
(B) Culex pipiens feeding on a seed pod.
(C) Anopheles sergentii feeding on a leaf in a blood meal-like fashion.
(D) Diagrammatic representation of the same bent labium for comparison to
the photograph.
14
Junnila et al.,2010
 analyses:
 Modified cold anthrone test for sugar,
 Cellulose particle staining for plant tissue,
 DNA sequencing for identifying plant sp.
15
150 males
and females
per each cage
Plant sugar meals of Anopheles sergentii in the lab:
Junnila et al.,2010
16
Tamarix nilotica
Ochradenus baccatus
Junnila et al.,2010
 192 field-caught An. sergentii were subjected to the
cold anthrone test- for fructose
 cellulose staining 200 field caught mosquito- for
plant tissue
 26 male and 26 female guts (n = 52) were extracted
and amplified by PCR- for what plant species
field-caught An. sergentii fed on
17
Plant sugar meals of Anopheles sergentii in the field:
Junnila et al.,2010
18
Conyza dioscoridis
Suaeda asphaltica
Malva nicaeensis Junnila et al.,2010
DECAYING FRUITS:
 Mosquitoes typically prefer damaged fruit over intact fruit
 Mosquitoes attracted to damaged Ceratonia siliqua
seed pods with fermenting liquid and to damaged
pomegranate are recorded in filed experiments
 Semiochemicals
 resources
 resource-dwelling microbes
Muller et al., 2011
19
ANT REGURGITATE:
 As a form of remarkable kleptoparasitism
 Cremastogaster spp-Malaya spp. interaction
 When a female mosquito inserts her proboscis into
the mouth of an ant, she induces trophallaxis and
 Then feeds upon the ant’s
regurgitate
Edwards, 1932
20
MOSQUITO POLLINATION:
21
•Among all pollinators, animals are estimated to be87.5%
•Within animals major pollinators are insects (around 90%)
•Hymenoptera and Lepidoptera are grab most of it.
•But Diptera are no less to it
•Mosquitoes are one that less know to us as a pollinators and
there role in ecosystem.
22
Thien, 1969
Mosquito pollination of Habenaria obtusata
23
Thien, 1969
OTHER PLANTS POLLINATED BY MOSQUITOES
 Spanish Catchfly, Silene otites is a flowering plant in the
family Caryophyllaceae
24
 mosquitoes also pollinate many Asteraceae plants
such as Goldenrods, Tansy plant etc.
Brantjes and Leemans, 1975
MOSQUITO POLLINATION:
25
Peach and Gries, 2019
26
Mean per cent seed set over different treatments
Peach and Gries, 2019
27
Number of mosquitoes field collected, and the abundance and
distribution of Asteraceae pollen present on the mosquitoes
Peach and Gries, 2019
28
Accumulation and distribution of pollen on Culex pipiens
mosquitoes after 2 h of experimental exposure to inflorescences
Peach and Gries, 2019
29
Peach and Gries, 2019
 Mosquitoes feeds on birds and mammals
 Even some mosquitoes feed on other insects
 Haematophagous insects can, and still do, feed on
larval stages of other insects
 Phytophagy Entomophagy Haematophagy
30
MOSQUITO FEEDING ON ANIMALS
31
Y-tube olfactometer
Determination of attraction of mosquitoes towards live
caterpillar or control using Y-tube
George et al., 2014
32
Per cent attraction of mosquitoes towards
live caterpillar and control
George et al., 2014
33
Y-tube olfactometer
Determination of attraction of mosquitoes towards live
caterpillar or dead caterpillar using Y-tube
George et al., 2014
34
Per cent attraction of mosquitoes towards live
caterpillar and dead caterpillar
George et al., 2014
35
Mean amount of CO2 released in live and dead
caterpillars of H. subfllexa and M. sexta
George et al., 2014
36
Determination of survival efficiency of mosquitoes on
the haemolymph of the caterpillars
George et al., 2014
37
Mean survival of mosquitoes when reared with and
without sugar water, and on caterpillar
George et al., 2014
BIZARRE CASES OF MOSQUITOES FEEDING ON
ANIMALS
 Annelids
 Earthworms
 leeches
 Amphibians
 toads
38
39
Female Uranotaenia sapphirina
Reeves et al., 2018
40
Females of Uranotaenia lowii
Reeves et al., 2018
HOW THESE MOSQUITOES LOCATE THEIR HOSTS?
 Multimodal cues guide them to their hosts
 Olfactory
 CO2
 Visual
 Thermal and etc.
 CO2 some times acts as sole cue or in combination
increases efficiency of the cue.
 Single cue
 Orienting towards artificial flowers(visual cue)
 Moving towards heat source having host’s temperature 41
42
Peach et al., 2019
Effect of olfactory and visual tansy inflorescence
cues on attraction of female Aedes aegypti and Culex
pipiens.
SL.NO. COMPOUND HUMAN SHARED?
1 butanoic acid Yes
2 2-methylpropionic acid Yes
3 2-methylbutanoic acid Yes
4 3-methylbutanoic acid Yes
5 benzoic acid Yes
6 hexanoic acid Yes
7 (−)-α-pinene Yes
8 −)-β-pinene NO
9 (−)-sabinene NO
10 (E/Z)-ocimene NO
11 germacrene-D NO
12 benzaldehyde Yes
13 acetophenone Yes
14 artemisia ketone NO
15 umbellulone NO
16 hexyl acetate NO 43
Headspace odorants of tansy inflorescences
Peach et al., 2019
FOSSIL EVIDENCE:
44
Culiseta species (Diptera: Culicidae), the fossil of a blood engorged
female mosquito in oil shale from northwestern Montana
Greenwalt et al., 2013
 As other organisms, mosquitoes also play role in
ecosystem such as pollination.
 Earlier, all mosquitoes are seems to be phytophagous.
 Mosquitoes use different array of resources, one among
them is human blood.
 Overlapping of cues between plants and vertebrates
supports the hypothesis that haematophagy may have
originated from phytophagy.
45
CONCLUSIONS:
46
Thank
you.

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Mosquito phytophagy, pollination and evolutionary transition to haematophagy

  • 1. MOSQUITO PHYTOPHAGY, POLLINATION AND EVOLUTIONARY TRANSITION TO HAEMATOPHAGY MADHUSUDAN M PALB-8138 Sr. M.Sc. Agril. Entomology 1
  • 2. PHYTOPHAGY: Phytophagy - feeding on the plant or/and plant derived products. DO FEMALE MOSQUITOES FEED ON PLANT/ PLANT DERIVED PRODUCTS? Yes 2
  • 3. DIFFERENT TYPES OF RESOURCES OF PLANTS  Floral nectar  Extra-floral nectar  Honey dew  Plant tissue  Ant regurgitates  Rotting fruits 3
  • 4. WHAT MAKES THEM(FEMALES) BLOOD FEEDERS? 4
  • 5. ARE THERE ANY ADVANTAGES OF SUGAR FEEDING?  Sugar feeding is a fundamental characteristic of mosquito life  Energetically, sugar and blood are interchangeable  The option to take sugar is retained for the competitive advantage  Even a replete blood meal does not initiate development if reserves become too low as a result of sugar deprivation  Sugar feeding enhances longevity and  Delayed egg laying (enhance fitness by being choosy) 5
  • 6. 6 Effect of sugar feeding on Anopheles gambiae on survival and biting frequencies.
  • 7. Water only Sugar + BloodBloodSugar solution 15151515 7 Mating X 5 Gary and Foster, 2001 Females only water Males 10% sucrose
  • 8. Males 10% sucrose Females only water Mating 8 Sugar solution Blood Sugar + Blood 1 X 20 cage 1 X 20 cage1 X 20 cage Gary and Foster, 2001
  • 9. Mean survival times of female Anopheles gambiae maintained on one of four diets 9 Age-specific survivorship (lx) of Anopheles gambiae females on four diets when females were either (A) pooled 15 to a cage or (B) isolated individually. Gary and Foster, 2001
  • 10. Proportion of Anopheles gambiae females biting each day, with access either to blood only or to blood + sugar 10 Gary and Foster, 2001
  • 11.  Plant source include both -carbohydrate (major) -non carbohydrate  Among all the sources- Floral nectar is the major source  But the content and nutrient status varies from source to source, and plant species to plant species 11 Are plants provide only sugar (CHO) to mosquitoes?
  • 12. 12 (A)Impatiens walleriana, (B) Asclepias curassavica, (C) Passiflora edulis, (D) Campsis radicans leaf, (E) C. radicans green seed-pod, (F) Ricinus communis. Positive control 10% sucrose in a 1.5 ml tube stuffed with cotton Negative controls 1. a drip irrigated soil pot, 2. a cotton-stuffed tube of water 3. an empty cage X 5 10 males + 10 females Effect of different nectar sources on survival of Aedes aegypti Chen and Kearney, 2015
  • 13. 13 days Proportion of mosquitoes surviving on different host plants and controls Chen and Kearney, 2015
  • 14. FEEDING ON PLANT TISSUE- IS IT A COMMON PHENOMENA? (A) Anopheles sergentii probing on a leaf. (B) Culex pipiens feeding on a seed pod. (C) Anopheles sergentii feeding on a leaf in a blood meal-like fashion. (D) Diagrammatic representation of the same bent labium for comparison to the photograph. 14 Junnila et al.,2010
  • 15.  analyses:  Modified cold anthrone test for sugar,  Cellulose particle staining for plant tissue,  DNA sequencing for identifying plant sp. 15 150 males and females per each cage Plant sugar meals of Anopheles sergentii in the lab: Junnila et al.,2010
  • 17.  192 field-caught An. sergentii were subjected to the cold anthrone test- for fructose  cellulose staining 200 field caught mosquito- for plant tissue  26 male and 26 female guts (n = 52) were extracted and amplified by PCR- for what plant species field-caught An. sergentii fed on 17 Plant sugar meals of Anopheles sergentii in the field: Junnila et al.,2010
  • 18. 18 Conyza dioscoridis Suaeda asphaltica Malva nicaeensis Junnila et al.,2010
  • 19. DECAYING FRUITS:  Mosquitoes typically prefer damaged fruit over intact fruit  Mosquitoes attracted to damaged Ceratonia siliqua seed pods with fermenting liquid and to damaged pomegranate are recorded in filed experiments  Semiochemicals  resources  resource-dwelling microbes Muller et al., 2011 19
  • 20. ANT REGURGITATE:  As a form of remarkable kleptoparasitism  Cremastogaster spp-Malaya spp. interaction  When a female mosquito inserts her proboscis into the mouth of an ant, she induces trophallaxis and  Then feeds upon the ant’s regurgitate Edwards, 1932 20
  • 21. MOSQUITO POLLINATION: 21 •Among all pollinators, animals are estimated to be87.5% •Within animals major pollinators are insects (around 90%) •Hymenoptera and Lepidoptera are grab most of it. •But Diptera are no less to it •Mosquitoes are one that less know to us as a pollinators and there role in ecosystem.
  • 22. 22 Thien, 1969 Mosquito pollination of Habenaria obtusata
  • 24. OTHER PLANTS POLLINATED BY MOSQUITOES  Spanish Catchfly, Silene otites is a flowering plant in the family Caryophyllaceae 24  mosquitoes also pollinate many Asteraceae plants such as Goldenrods, Tansy plant etc. Brantjes and Leemans, 1975
  • 26. 26 Mean per cent seed set over different treatments Peach and Gries, 2019
  • 27. 27 Number of mosquitoes field collected, and the abundance and distribution of Asteraceae pollen present on the mosquitoes Peach and Gries, 2019
  • 28. 28 Accumulation and distribution of pollen on Culex pipiens mosquitoes after 2 h of experimental exposure to inflorescences Peach and Gries, 2019
  • 30.  Mosquitoes feeds on birds and mammals  Even some mosquitoes feed on other insects  Haematophagous insects can, and still do, feed on larval stages of other insects  Phytophagy Entomophagy Haematophagy 30 MOSQUITO FEEDING ON ANIMALS
  • 31. 31 Y-tube olfactometer Determination of attraction of mosquitoes towards live caterpillar or control using Y-tube George et al., 2014
  • 32. 32 Per cent attraction of mosquitoes towards live caterpillar and control George et al., 2014
  • 33. 33 Y-tube olfactometer Determination of attraction of mosquitoes towards live caterpillar or dead caterpillar using Y-tube George et al., 2014
  • 34. 34 Per cent attraction of mosquitoes towards live caterpillar and dead caterpillar George et al., 2014
  • 35. 35 Mean amount of CO2 released in live and dead caterpillars of H. subfllexa and M. sexta George et al., 2014
  • 36. 36 Determination of survival efficiency of mosquitoes on the haemolymph of the caterpillars George et al., 2014
  • 37. 37 Mean survival of mosquitoes when reared with and without sugar water, and on caterpillar George et al., 2014
  • 38. BIZARRE CASES OF MOSQUITOES FEEDING ON ANIMALS  Annelids  Earthworms  leeches  Amphibians  toads 38
  • 40. 40 Females of Uranotaenia lowii Reeves et al., 2018
  • 41. HOW THESE MOSQUITOES LOCATE THEIR HOSTS?  Multimodal cues guide them to their hosts  Olfactory  CO2  Visual  Thermal and etc.  CO2 some times acts as sole cue or in combination increases efficiency of the cue.  Single cue  Orienting towards artificial flowers(visual cue)  Moving towards heat source having host’s temperature 41
  • 42. 42 Peach et al., 2019 Effect of olfactory and visual tansy inflorescence cues on attraction of female Aedes aegypti and Culex pipiens.
  • 43. SL.NO. COMPOUND HUMAN SHARED? 1 butanoic acid Yes 2 2-methylpropionic acid Yes 3 2-methylbutanoic acid Yes 4 3-methylbutanoic acid Yes 5 benzoic acid Yes 6 hexanoic acid Yes 7 (−)-α-pinene Yes 8 −)-β-pinene NO 9 (−)-sabinene NO 10 (E/Z)-ocimene NO 11 germacrene-D NO 12 benzaldehyde Yes 13 acetophenone Yes 14 artemisia ketone NO 15 umbellulone NO 16 hexyl acetate NO 43 Headspace odorants of tansy inflorescences Peach et al., 2019
  • 44. FOSSIL EVIDENCE: 44 Culiseta species (Diptera: Culicidae), the fossil of a blood engorged female mosquito in oil shale from northwestern Montana Greenwalt et al., 2013
  • 45.  As other organisms, mosquitoes also play role in ecosystem such as pollination.  Earlier, all mosquitoes are seems to be phytophagous.  Mosquitoes use different array of resources, one among them is human blood.  Overlapping of cues between plants and vertebrates supports the hypothesis that haematophagy may have originated from phytophagy. 45 CONCLUSIONS:

Editor's Notes

  1. Males placed in an 85-liter acrylic cage with water and 10% sucrose soaked cotton wicks. The water was replenished as needed, and the sucrose was replaced every 5 d. The females were placed in a small acrylic plastic cage (15 by 21 by 27 cm) supplied with water only. Three days after eclosion, the females were placed in the large male cage for free mating overnight. Sucrose was absent from the male cage during this time. After 24 h with males, 15 females were placed into each of four small acrylic cages Cages of mosquitoes receiving blood were given the opportunity to feed daily in the morning on the hand and forearm of a human host (R.E.G.) for 10 min.
  2. Vertical arrows point to the median survival time of females on each diet.
  3. Impatiens More attractive and helped mosquitoes to survive for longer time provide evidence that they give more than just sugar.
  4. Fresh branches, including green foliage and flowers, of Tamarix niloicata or Ochradenus baccatus were presented to adults in separate cages. Mosquitoes were killed with CO2 in the following morning
  5. Of these, 22 were PCR positive for the rbcL fragment (Table 2). On sequencing, 15 guts (6 males and 9 females) contained the rbcL sequence of S. asphaltica, 5 guts (all males) contained the rbcL sequence of Malva and 2 guts (1 male and 1 female) contained the sequence of C. dioscoridis. From 96 tested male guts, 24 were sugar positive and from 96 tested female guts, 29 were sugar positive. From 100 dissected male guts, 32 contained stained cellulose particles From 100 dissected female guts, 38 contained stained cellulose particles.
  6. Repeated encounters and mutual disturbance of Hodgesia mosquitoes and ants at damaged-plant-tissue feeding sites, or mosquito consumption of honeydew in the presence of ants, may indicate events or circumstances that have given rise to the evolution of this form of kleptoparasitism.
  7. , aFor the cross-pollination treatment, we kept a mixed-sex group of 60 mosquitoes without water or sugar water for 3 days in a pollinator exclusion bag (3.78 L), enclosing one experimental inflorescence of a living (potted) greenhouse-grown plant and two inflorescences excised from a different plant, retaining each cut inflorescence in a water-filled, 20-mL vial. The control treatment was identical except that mosquitoes were absent from the pollinator exclusion bag. pollinator exclusion bag
  8. where the cross- and self-pollination treatments yielded a total of 394 and 99 seeds, respectively, whereas the control treatment yielded only 13 seeds. Mosquito-mediated self-pollinated inflorescences (n = 13) and cross-pollinated inflorescences (n = 12) had a mean successful seed set per inflorescence of 0.22 % (95 % CI 0–0.61 %) and 0.97 % (95 % CI 0.57–1.37 %), respectively. Control inflorescences (n = 16) without exposure to mosquitoes had a mean successful seed set per inflorescence of 0.018 % ex 2 mosquito-mediated cross-pollinated inflorescences (mean: 12.5 %; control inflorescences (mean: 0.06 %; The cross-pollination treatment (n = 11) yielded a combined total of 3939 seeds, whereas the control treatment (n = 10) yielded a total of only 14
  9. We ran sampling surveys during the scotopic periods of (1) 00:00–01:00 h on each of 31 July and 04August, (2) 22:30–23:00 h on each of 11 and 18 August, and (3) 21:00–22:00 h on 22 August. Illuminating plants with a headlight, we aspirated mosquitoes while they were probing for nectar and then placed each mosquito in a separate sealable sandwich bag, Achillea millefolium
  10. we placed a mixed-sex group of 60 Cx. pipiens in a mesh cage (30 9 30 9 46 cm high) that enclosed field-collected inflorescences of either T. vulgare or A. millefolium or inflorescence of S. canadensis. Exposed mosquitoes for 2 hr.
  11. male Culex tarsalis, female Cx. Pipiens from Achillea millefolium, male and female Culex pipiens feeding on inflorescences of greenhouse-grown Tanacetum vulgare d pollen on the antennae of a feeding female female with numerous pollen grains on her proboscis, palps, antennae, legs, and thorax, f. using her forelegs to remove pollen from her proboscis
  12. they have not completely abandoned the entomophagous behavior during evolution and it is still conserved as an opportunistic behavior among dipteran blood feeding insects such as mosquitoes.
  13. Four-to-six day old lab-reared Female An. stephensi were released at the main, downwind end of the Y-tube with charcoal-purified air pushed through both arms of the Y-tube at the rate of 0.75 litre/sec. One 4th-instar laboratoryreared larva of either H. subflexa or M. sexta was placed on a cardboard disc and retained inside the arms of the Y-tube at the extreme upwind end. Mosquitoes were assayed individually and those that flew upwind the length of the tube and landed on or near the caterpillars or in the control arm (cotton dental wick of the same dimension caterpillar)
  14. Mosquito took only less than 10 seconds to enter the y-tube arm containing the caterpillar and only 15–20 seconds to land on the caterpillar attracted to live M. sexta (87%) compared to the clean air cotton wick control arm (13%) H. subflexa caterpillars, 82% 18%
  15. Choice assays were performed using live caterpillars vs. a blank (dental wick) control, and in addition, live caterpillars were compared against dead caterpillars that had been frozen for 12 hours at 220uC, and then thawed back to room temperature.
  16. H. subflexa, 81% of the females flew to the arm containing live caterpillars compared to only 19% flying to the dead ones. M. sexta caterpillars, 79% of females flew upwind to the live caterpillars compared to only 21% to the dead
  17. Carbon dioxide Measurement using Respirometer Assays: Metabolic rates were measured by placing each live or dead caterpillar into its individual flow-through respirometry chamber result - live caterpillars emitted ca. a 10- times greater amount of CO2 than did dead caterpillars Thus CO2 emissions from the caterpillars could possibly have contributed to the elevated levels of attraction of An. Stephensi females
  18. Female An. stephensi that were four to six days The mosquitoes were never blood-fed, and were given access to 10% sugar solution until the start of the experiment. Forty mosquitoes were introduced into a rectangular plastic cage The experiment consisted of three treatments and four replicates for each treatment (N= 160). 10% glucose solution in a 40 ml glass bottle with filter paper wicking the sugar solution In the second treatment, three 4th -instar H. subflexa larvae were placed inside a second cage and allowed to roam around the cage. 3 no food no water.
  19. Only 40% of the females caged in the presence of caterpillars were dead by Day 4 compared to 85% of those caged with no food. By day 5, mortality of females in the no food cages had increased to 98%, compared with 60% in cages containing caterpillars. Females provided with continuous access to sucrose water survived the longest, exhibiting only 20% mortality after 10 days.
  20. Trap captures of 1- to 3-day-old female A. aegypti and 1- to 3-day-old female C. pipiens in response to visual and olfactory tansy inflorescence cues. A rectangular box with hatched lines indicates that the occluded inflorescence offered no visual cues. numbers within bars indicate the mean percentage of mosquitoes not captured. stem of inflorescence and occluded with 3 layer of cheese cloth with proper mess size which allow volatiles to pass
  21. Overlapping cues between plants and vertebrates support the postulated concept that haematophagy of mosquitoes may have arisen from phytophagy. Porapak-Q™ adsorbent
  22. The existence of this rare specimen extends the existence of blood-feeding behavior in this family of insects 46 million years into the past. Heme, the oxygen-carrying group of hemoglobin in the host’s blood, was identified in the abdomen of the fossil mosquito by nondestructive mass-spectrometry analysis. Although large and fragile molecules such as DNA cannot survive fossilization, other complex organic molecules, in this case iron-stabilized heme, can survive intact and provide information relative to the mechanisms of the fossilization process.