Longevity March 2022a
Longevity March 2022a
Longevity March 2022a
com
Guppy Longevity
by Diana Walstad (March 2022)1
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This article complements two other articles: ‘Small-Scale Guppy Breeding’ and ‘Breeding Guppies: Genetic
Pitfalls and Successes’ that are also available on my website: http://dianawalstad.com
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Lifespan of Guppies
In 1961, the British scientist
Comfort [1] officially reported a
potential lifespan of 5 years for the
guppy Poecilia reticulata.
For his comprehensive longevity
study [2], he used 4 domestically
bred strains. Young guppies were
sexed and placed in containers with
gravel, snails, and floating plants
until their death.
Apparently, he provided the
most basic of care, ”Aquaria looked
most unsatisfactory, but the fish in
them lived for 4-5 years, as against a
normal aquarium life of 2 years or
less.” No filters or aerators. They
were fed live Tubifex worms once a
week and a cereal supplement.
Water was changed only when the
fish showed “discomfort.” Fig 3. Survival of 1950s Guppies [2]
Results are for 312 males and 351 females kept in 50-liter (~13
Fig 3 shows results from his
gal) single-sex tanks until their death. Each tank held either 12-20
longevity study. Fish started out virgin females or 20-35 virgin males. Fish were counted for
with 100% survival when they were survival at 100-day intervals. I show here only his tank results, but
100 days old. At 2 years, survival survival was similar for guppies kept in jars.
of both males and females had {I drew graph from tabulated data in Comfort’s paper.}
decreased to 55% - 60% (See
graph’s vertical bar). Males began
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dying slightly sooner than females. At 1500 days (4.1 years), all males were dead, but 12% of females
were still alive. The last females died at 1900 days (5.2 years) of age. The investigator noted that
disease (e.g., ‘Fish TB’) and other pathologies increased with aging.
Comfort’s survival data applies to virgin guppies, which have a longer lifespan than reproducing
guppies [3]. Nevertheless, the 5-year lifespan assigned by Comfort to the species [1] is amazing
compared to the brief life of today’s fancy guppies.
Wild guppies also have a notably greater longevity than today’s domestic guppies. In a
comprehensive study,
Reznick [4]
documented the
longevity of recent
descendants of wild
guppies. Average
longevity for
reproducing females
was 2.3 years.2
Another
investigator (Evans)
[6] conducted an in-
depth analysis of
fecundity (or fertility)
in female feral
guppies. Evans’ data
also provided me with
information on female
longevity, for the fact
is that only live
females reproduce.
The study’s 84
virgin females were
mated (at age 3
Fig 4. Longevity and Fecundity of Feral Guppies [6]
months) to 1-2 males
Columns shows the average brood size (± S.E.) for the study’s 81 females over
during each of their 18 sequential brood cycles. The number of females still producing for each cycle
brood cycles. Fig 4 (‘n’) are shown in the columns.
shows the fecundity {I drew graph from raw data provided by the investigator (Evans) in 2020.}
of the 81 reproducing
females (3 of the 84 females produced no offspring.). First broods averaged only ~5 fry for 81 females.
By the 3rd brood cycle, the average number of offspring had tripled to over 15 for 77 females. By the
15th cycle, the average reached a peak at ~31 fry for the 10 remaining females. At the final 18th cycle,
2
Wild guppies were captured from two collection sites differing in predation level within two different river
systems. F2 females produced from each of the 4 populations were maintained on either a high or a low feeding
regimen to produce 8 groups (n = ~28 fish/group). The 226 females were isolated at 25 days in 7.8 liter
containers. They were mated throughout their lives, first as virgins and then after each parturition [4, 5]. The
average lifespan for all 8 groups (a sampling distribution) was 846 days (± 42 S.E.) or 2.3 years. The range for
individual lifespan was 151 - 1,464 days.
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the 3 still-reproducing females produced an average of only 5 offspring; their reduced fecundity was
probably due to aging ovaries.
Interestingly, a huge surge in fecundity followed the 9th brood cycle. I think that the weaker
females had died off and the remaining, more robust females had come into their own. They were older,
and thus, had larger broods.
Statistical summations can be misleading. The 84 females produced an average of 7.6 broods per
female, yet many individuals produced far more. For example, 21 females (or 25% of the starting
population) produced at least 11 broods. The graph (Fig 4) presents a far more realistic picture of
female longevity and reproductive fitness than population averages.
Predicting Longevity
Evans’ study [6] also provided me with data on individual females so that I could answer the
following question, “Could I distinguish superior females from weak females based on their
reproductive performance while young?
To answer this question, I picked 8 females on each end of the longevity/reproductive fitness
spectrum shown in Fig 4. My 8
“weakest” females produced only 4 Number of Offspring Weakest Fittest
broods (probably due to death) Females Females
during their short reproductive First Brood 6.6 (±3.9) 5.1 (±3.6)
lifespan. In contrast, my 8 “fittest”
females produced ≥ 16 broods. Second Brood 13 (±3.6) 12 (±5.2)
Assuming a typical one-month Third Brood 17 (±3.2) 18 (±5.1)
interval between mating and broods,
the 8 fittest females would have lived Fourth Brood 18 (±5.6) 13 (±10)
at least 20 months (3 + 1 + 16 = 20). Lifetime Production
Did the 8 weak females signal 55 (±10) 267 (±40)
their poor future early in life? The Table 1. Fitness Extremes in a Population [6]
results in Table 1 say, “No.” Numbers are the average (± S.D.) of offspring per brood. The
Average size of the first 4 broods of ‘Weakest Females are the 8 females that produced only 4 broods
the two female groups (“weakest” during their lifespan. The 8 ‘Fittest’ females produced ≥ 16 broods.
{I constructed table from raw data provided by Evans in 2020.}
and ‘fittest”) was very similar.
Indeed, the weakest females actually
produced more offspring than the fittest females (18 v. 13) for their fourth and last brood. Because the
fittest females continued reproducing long after their first 4 broods, their life-time output was over 4
times greater than that of the weakest females (267 v. 55 offspring). The fittest females not only lived
longer but were more productive.
Results from Table 1 support my own observation that future longevity and reproductive success in
young fish is difficult or well-nigh impossible to predict. Indeed, when I examine my own young
guppies of 4 months age, I have no idea which ones will outlive their siblings. Only by waiting does the
inherent weakness of some individuals—and the superiority of others—become apparent.
salt) to prevent disease outbreaks. Should an outbreak occur, they may be treated for the disease. Over
multiple generations, domestication inevitably results in lowered immunity.3
Many guppy breeders select primarily for color, fin shape, and strain uniformity. Longevity is rarely
considered. Indeed, when my male BG guppies died at ~5 months, I simply replaced them with younger
males. Only later, did I begin to contemplate the negative consequences of this practice.
For guppies, tail or caudal size seems to be another factor affecting longevity, with swordtail (short-
tail) strains living longer than delta tail strains. The consensus among most guppy breeders is that delta
strains have a lifespan of 1-1.5 years; swordtail strains, 1.5-2 years [8]. Because of their large heavy
tails, older delta males are purportedly less able to inseminate females. In 1961, a master guppy breeder
Paul Hahnel [9] recommended using younger males as breeders. No doubt, many breeders maintaining
delta strains have followed his advice.
Guppy breeders and scientists [10] often cite inbreeding as a cause of reduced longevity. Perhaps,
but I believe there are two other causes that get much less attention.
3
One investigator [7] showed that wild guppies became much more susceptible to gyrodactylid parasites (i.e.,
skin flukes) following just 4 generations of lab maintenance. The survival rate declined from 96% to 58% due to
domestication.
4
The rate for loss of genetic variation is 1/(2NE) per generation interval (G) [11]. (NE is the ‘Effective Population
Size’ or the number of breeding individuals.)
5
Similarly in Drosophila lab strains, the use of only younger flies for breeding reduces fly longevity [13]. Most
Drosophila research labs maintain their strains with a strict 14-day generation interval, despite the fact that wild
flies can live ~80 days [14]. One study [13] found that male life expectancy was only 60 days for lab flies
compared to 83 days for wild flies. Other investigators [15] reported that the average lifespan for reproducing
females from a wild population decreased from 44 to 31 days after two years of lab maintenance.
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One German investigator [18] estimated that his 50-year-old strains had undergone 200 generations
in captivity. That translates into an average generation interval of only 4 months (200 ÷ 50 = 4). He
also reported that many of his females died before they produced their third brood.
Many scientists have observed a trade-off between early reproduction and longevity [14]. It
supports a widely accepted theory for the evolution of aging.6 Individuals that invest more in early
reproduction do so at the expense of their body maintenance, resulting in a shorter lifespan. So, if you
use only young guppies for breeding, you are in essence selecting for early maturity at the expense of
longevity.
Virgin Guppies
Almost all serious guppy breeders and scientists work with virgin females. That way, they know
exactly which male is the sire of a particular female’s offspring. (While the most recent mating
generally gets paternity precedence, one is never quite sure who is the sire when dealing with non-virgin
females. Also, you may get mixed batches sired by multiple males.)
Working exclusively with virgins, though, can decrease female longevity. Virgins get little exposure
to the stresses of mating and reproduction. In many show breeding setups, a female may only produce 3
litters during her entire life, whereas females that have grown up with their male siblings will reproduce
continuously throughout their lives. For example, wild guppies average 22 litters per female.7
Thus, there is almost no selection for genes that would provide females with resistance to mating
injuries and the normal stress of giving birth month after month. It is not surprising that hobbyist report
a litany of problems in their female guppies.
Longevity v. Reproduction
In breeding guppies for increased longevity, reproductive lifespan may be just as important as
longevity. In breeding my own guppies for longevity, I came across males and females that lived longer
but had stopped reproducing months earlier. Fig 6 shows a 15-month-old male that I was unable to get
any progeny from. When he was a year old, I put him with females for 3 months. The females
produced progeny, but the batches were all sired by earlier males.
Guppy breeders should be aware that there is a known trade-off between reproduction and longevity.
If one focuses solely on longevity, the result may be long-
living guppies that cannot reproduce.
Wild female guppies were found [4] to have an
average lifespan of 2.3 years and a reproductive lifespan
of 2.0 years representing 87% of their total lifespan.
However, these population averages encompass
enormous individual differences.10
Indeed, Reznick [5] reported that reproduction was
highly variable in his females. Some females produced a
new litter every month for years and then died promptly
after their last litter; others skipped litters for months
before getting back on track; some lived many months
after their last litter.
There are no similar comprehensive guppy studies on Fig 6 Long-living but Infertile
males. Based on work with Drosophila, however, I Unfortunately, I was unable to get any
progeny from this beautiful male as an older
suspect that male guppies have a longer reproductive
adult (~12 months). Photo shows him at 15
lifespan than females. Male fruit flies are fertile months.
throughout their lives, while female flies may live long
after they have stopped reproducing [14].
10
The average reproductive lifespan (i.e., ‘age of last reproduction’) for the 8 groups (See Footnote #2) was 727
days (± 43 S.E.) or 2.0 years. The range for individuals across the groups was 109 - 1,294 days [4].
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In keeping older guppies until they run Q. If I use older females for breeders (select for greater
out their internal clocks, two cautions: female longevity), will that also increase male longevity?
First, as a fish ages, it becomes
increasingly vulnerable to endemic, potential A. It could, especially if your guppies are dying at a very
pathogens and disease. If an older fish—or young age (5-6 months). Some strains of fancy guppies
any fish for that matter—becomes a disease are so genetically weak that almost any counter-measure
reservoir, it can endanger tankmates. Thus, I would probably help. Increasing immunity to bacterial
monitor the health of older fish a little more infections certainly would benefit both sexes.
closely than younger ones. Should one Beyond a certain point, though, achieving greater
longevity in one sex does not guarantee longevity in the
develop disease symptoms or become
opposite sex.
incapacitated, I euthanize it in a timely
manner.
Second, animals are known to acquire deleterious mutations in the germline as they age.11 (They
become increasingly less able to repair the precious DNA within their reproductive organs.) Thus, when
a very old female guppy is no longer reproducing normally with her usual large batches, I would not
save progeny from her last small batches. Reduced fecundity due to ageing correlates with deleterious
mutations in the fry’s own germline DNA. Those mutations will be inherited by the next generation.
Readers should understand that my experience with guppy longevity to date (2022) is limited.
Breeding guppies for longevity is a step-wise process that takes time. However, I enjoy having beautiful
guppies that stick around a little longer than a few months.
❖ Save progeny from older—but not too old! —guppies. (Older individuals have proved their
superiority by outliving their siblings.)
❖ Recognize that increasing longevity may require selection over multiple generations.
❖ Keep in mind that older reproducing females have much larger litters than young females.
This increases your opportunities for finding superior individuals.
❖ Consider outcrossing to strains that are recognized for greater longevity.
❖ Avoid severe inbreeding (e.g., mating full-siblings) over multiple generations. Instead, plan
matings between less related guppies (e.g., cousins and half-siblings).
❖ Be judicious in treating guppies for disease. Concentrate more on breeding disease-resistant
fish. Use disease outbreaks (e.g., flukes and Costia) to cull susceptible individuals lacking
genes for normal immunity and select out more resistant individuals.
11
The ‘Lansing Effect’ is a well-known problem in human fertility clinics. For example, children born to aged
parents have an increased risk for inheriting genetic disorders (e.g., autism and schizophrenia). However in
guppies, saving progeny from older individuals may be less risky. Thus, in an experimental study with feral
guppies, Gasparini [23] showed virtually identical sperm quality and siring success for older males (age 14-
months) as compared to younger males (age 5-months).
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References
1. Comfort, Alex. 1961. The lifespan of animals. Scientific American 205: 108-119.
2. Comfort, Alex. 1961. The longevity and mortality of a fish (Lebistes reticulatus Peters) in captivity.
Gerontologia 5: 209-222.
3. Reznick DN. 1997. Life history evolution in guppies (Poecilia reticulata): guppies as a model for studying
the evolutionary biology of aging. Experimental Gerontology 32: 245-58.
4. Reznick DN et al. 2004. Effect of extrinsic mortality on the evolution of senescence in guppies. Nature 431:
1095-99.
5. Reznick D et al. 2006. The evolution of senescence and post-reproductive lifespan in guppies (Poecilia
reticulata). PLOS Biology 4: 136-143.
6. Evans JP. 2012. Lifetime number of mates interacts with female age to determine reproductive success in
female guppies. PLOS ONE 7: e47507 (6 pp.)
7. van Oosterhout C et al. 2007. The guppy as a conservation model: implications of parasitism and inbreeding
for reintroduction success. Conservation Biology 21: 1573-83.
8. Discussion (October 2018) about longevity on the Guppy Gene Collectors forum
https://www.facebook.com/groups/GuppyGeneCollectors/
9. Whitney LF. and Paul Hahnel. 1964. All About Guppies. T.F.H. Publications (Neptune City, NJ), 128 pp.
(Authors’ advice on using younger males for breeding is on page 87.)
10. Chiyokubo T et al. 1998. Genetic features of salinity tolerance in wild and domestic guppies (Poecilia
reticulata). Aquaculture 167: 339-348.
11. Allendorf FW et al. 2013. Conservation and the Genetics of Populations (2nd Edition) Wiley-
Blackwell Publishers (West Sussex, UK), p 131.
12. Bias, Alan S. Dec. 25, 2011a. Breeding Strategies & Genetic Consequences in Guppies. (PDF of 11 pages
can be downloaded from: https://independent.academia.edu/AlanBias)
13. Linnen C, Tatar M and D Promislow. 2001. Cultural artifacts: a comparison of senescence in natural,
laboratory-adapted and artificially selected lines of Drosophila melanogaster. Evol Ecol Res 3: 877-88.
14. Flatt T. 2020. Life-history evolution and the genetics of fitness components in Drosophila melanogaster.
Genetics 214: 3-48.
15. Sgro CM and L Partridge. 2000. Evolutionary responses of the life history of wild-caught Drosophila
melanogaster to two standard methods of laboratory culture. American Naturalist 156: 341-353.
16. Reznick DN et al. 2001. The evolution of senescence in natural populations of guppies (Poecilia reticulata):
a comparative approach. Experimental Gerontology 36: 791-812.
17. Larsen LK et al. 2011. Temporal change in inbreeding depression in life-history traits in captive populations
of guppy (Poecilia reticulata): evidence for purging? J Evol Biol. 24: 823-34.
18. Farr JA. 1980. Biased sex ratios in laboratory strains of guppies, Poecilia reticulata. Heredity 47: 237-248.
19. Zwaan B et al. 1995. Artificial selection for developmental time in Drosophila melanogaster in relation to
the evolution of aging: direct and correlated responses. Evolution 49: 635-648.
20. Kwan L et al. 2013. Sexual conflict and the function of genitalic claws in guppies (Poecilia reticulata). Biol
Lett 9: 20130267.
21. Greven H. 2005. Structural and behavioral traits associated with sperm transfer in Poeciliinae. In: Grier HJ
et al, Viviparous Fishes: Proceedings of the I and II International Symposia on Livebearing Fishes, New Life
Publications (Florida), pp 147-165.
22. Parker GA and L Partridge. 1998. Sexual conflict and speciation. Phil Trans R Soc B 353: 261-274.
23. Gasparini C et al. 2010. Effect of male age on sperm traits and sperm competition success in the
guppy (Poecillia reticulata). J. Evol. Biol. 23:124-135.
Diana Walstad is the author of Ecology of the Planted Aquarium (2013). For articles
and more information about her books, see: http://dianawalstad.com