Longevity March 2022a

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Guppy Longevity
by Diana Walstad (March 2022)1

Guppies are no longer the hardy fish they once were.


1950s guppies reportedly had a 2-5 year life span [1]. Q&A I’ve never been successful with guppies
Initially, I blamed premature death on the seismic in spite of every aquarium article saying they’re
shift from “basement bred” American guppies to mass- easy, so I finally gave up my serial killer ways
and stopped buying the poor things. I’m now
produced guppies from Southeast Asia. However,
wondering if it wasn’t me at all.......
research facilities also report a surprisingly short lifespan
in their established lab strains. Answer Don't blame yourself. No matter how
Granted, environmental factors (temperature, food, good the care, many domesticated guppies have
water quality, etc.) will affect lifespan. However, I focus a lifespan shorter than the two years that it
here on the genetic causes. Many common guppy could be. For they are genetically weak. They
breeding practices—in the home and in research labs— have little disease resistance and are
inadvertently result in an inherited short lifespan. physiologically unfit. They may thrive in the
To simplify a complex subject, I describe experiences original breeder’s tanks or ponds, but they have
with my own guppies. I was able to increase the lifespan trouble when forced to adapt to a new
of my guppies from about 6 months in 2017 to 12 months environment—your tank.
in 2022—a doubling of the generation interval. Since Many standard guppy breeding practices
inadvertently produce genetically weak fish.
guppies can live two years or more, it is admittedly a
modest achievement with more work ahead.
However, I did not select solely for longevity. For what good is a long lifespan if the resulting
guppies are unattractive and don’t reproduce? My goal is to have pleasing, attractive, normally
reproducing guppies that also have a decent lifespan.

Short Life in a Blue Grass Strain

Longevity in guppies became an issue when I


started up again with guppies in 2017. After a
long hiatus with other aquarium fish (Rainbowfish,
cichlids, etc.), I returned to my first love—guppies.
I purchased different fish from a variety of
sources and strains. The BG (Blue Grass) strain of
guppies (Fig 1), two pairs purchased via the
Internet, were the most pleasing. They were big,
Fig 1. My Blue Grass (BG) Strain of guppies,
beautiful, prolific, and relatively disease-resistant.
purchased in 2017, was almost—but not quite—
It took me almost a year to realize that these perfect. In my tanks, the fish did not live much longer
gorgeous guppies were just not living that long. than 6 months. (The original breeder reported a
In 2018 I started monitoring their longevity. In lifespan of about 8 months in his tanks.)
July, a prized BG female produced a large batch.
Raised outside in summer tubs and exposed to the elements, the offspring were uniformly vigorous.
When they were 3 months old, I culled small unattractive individuals, sold some, and retained 2 males
and 7 females for breeding. I then documented the lifespan of the 9 select individuals.

1
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
2

Unsurprisingly, the males did not live


long—about 4-5 months. Irreversible decline
began when they rested on the bottom with
swollen bellies and lost all interest in food and
females. As to the females… From my 7
select females, I had expected to obtain several
good breeders. Alas, I ended up with only
Fig 2. Female #3 (r) at Five Months Age had one—Female #3. At 5 months age, her
grown noticeably larger than her 6 sisters. She outlived superiority became evident (Fig 2).
them all. Before dying at 9 months, due to a prolapsed I believe that the short lifespan of my BG
uterus, she produced several large batches from which I strain was caused by genetic “plumbing
selected the next generation. She was the foundation problems.” The males suffered from intestinal
female for my longevity project. blockage; the females, reproductive problems.
The short lifespan of this strain—and
probably many other fancy domestic strains—is not unusual. It contrasts markedly with the natural
lifespan of the species.

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
3

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.
4

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.

Breeding Practices that Shorten Longevity


Domestication inevitably leads to some loss of genes for longevity. Domesticated guppies are fully
fed, never preyed upon, and kept in pristine water. Sometimes, the water contains prophylactics (e.g.,
5

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.

Short Generation Intervals


Generation interval (G) affects genetic health just as much as the size of the breeding population
[11].4 As population longevity and G (the time between each generation) increases, the rate loss of
genetic variation and the accumulation of deleterious mutations automatically slow down.
When only young fish are used for breeding, longevity and G gradually erode. Ever wonder why
show strains lack the longevity of wild populations? Most show breeders use only the first few batches
from their females. Once the females have produced about 3-4 batches, the females are discarded or no
longer used for breeding [12].
When G is shortened artificially in this manner, it leads to all kinds of genetic troubles including
reduced longevity.5 Weak females get the same chance to pass on their genes as superior females. Ditto
for the males. There is no selection for longevity, reproductive fitness, disease resistance, etc.
In nature, a fish’s life may also be cut short abruptly, but weaker fish—whether old or young—
generally succumb first allowing fitter fish to continue reproducing. Older fish still in their prime are
not arbitrarily removed. Moreover, older females produce more offspring than younger females [16].
Thus, a wild population will contain a greater proportion of progeny from fitter females.
Norwegian investigators [17] documented a gradual shortening of wild guppy lifespan over the
course of 10 generations of lab breeding. Each generation was started with progeny from only the
female’s first 1-3 broods. (Females were given three chances to produce a brood of at least 5 fry.)
Thus, there was no selection for females that could produce more than 3 broods. At the start, a full 97%
of the guppies survived to 12 months. After 10 generations, however, survival declined to 87%.
Apparently, a decrease like this in scientific labs is not uncommon.

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.
6

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.

Effect of Reproduction on Longevity


Mating and reproduction decreases longevity. Reznick [3] estimated that the “half-life” of Comfort’s
virgin females was ~40 months compared to only ~14 months for his (i.e., Reznick’s) reproducing
females.8
For females, the costs of mating and reproduction are particularly severe.9 For example, the male
gonopodium has a “claw” that grips inside the female’s reproductive tract during copulation. This claw
prolongs copulation, allowing the male to transfer more sperm during non-consensual sex [19], but it
can tear female tissue [21]. This tearing allows the entry of foreign bacteria. If the female has inherited
good immunity, she may be okay, if not, she could develop a widespread and deadly internal infection.
In Drosophila and many other taxa, the male ejaculate contains various seminal fluid proteins
designed to increase a male’s mating success. Some proteins may decrease a female’s receptivity for
remating. Others may disable competing sperm and increase egg production. Inadvertently, some
6
The association of early reproduction with a short lifespan is a classic example of antagonistic pleiotropy (AP).
Some genes exert benefits to a species in early development (increased fecundity) but bad effects later in life
(early senescence). The AP theory of aging explains why senescence, a deleterious condition, is not selected out
during evolution.
7
The 22 (± 1.7 S.E.) litters/female represents a sampling distribution for 8 diverse groups of wild guppies. The
range for individuals across the groups was 2-43 litters/female [4]. (See Footnote #2 for a more complete
description of the groups studied.)
8
Drosophila studies have consistently shown that mating reduces the fruit fly’s lifespan, particularly in females
[14]. Zwaan [19] reported 28% and 44% mating-related decreases for males and females, respectively.
9
Males pay less of a price for reproduction. Continuous courting and competition between males exert a toll, but
the cost (in terms of lifespan) is much less than for females.
7

proteins have side effects that are known to decrease


female longevity [22]. If a female in a given population
has resistance to these proteins, she may be okay; if not,
her life may be cut short.
No doubt there are many other reproductive factors
that can shorten a female’s lifespan. In 2017 after I
restarted with guppies, I witnessed the distress and death
of several females shortly after they gave birth [Fig 5].
Females have developed a range of protective
mechanisms that counter male strategies to increase Fig 5 Reproduction is Stressful
paternity [22]. Those mechanisms inevitably involve Female #3 (See Fig 2) had out-lived all
inheritance, evolution, and selection for genes that her siblings. But she became morbid at 9
months, a few days after parturition. Note
enhance a female’s reproductive fitness. But when there
swelling at vent area, possibly due to a
is little selection for females that can handle the natural prolapsed uterus. I blame this on a weak
stresses of reproduction, those genes can be lost. uterus.

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].
8

Breeding Older Fish


Older guppies are valuable as breeders simply because they
have proven their ability to survive. Moreover, older females
produce much larger broods. A female’s first brood might number
less than 10 fry, but a large older female (7-12 months) can
produce more than 60. That gives the guppy breeder more
progeny to choose from, increasing the odds of finding individuals
that are both attractive, sturdy, reproduce normally, and that have
longevity.
At about 3-4 months age, I select individuals that I find
personally pleasing and put them in a separate tank. I want
females with good size, nice shape, and big dorsals. For males, I
Fig 7 The “Fin Rot” shown in
want big spotted dorsals, short backs, energy, lots of color, etc. this otherwise healthy male only
After this initial selection, I may do some miscellaneous became apparent at an older age (7
culling. As the guppies approach 8 months age, markers for months).
longevity and fitness become increasingly apparent. By then
some individuals have died out; some have gained size on the others. Some males have lost their energy
and enthusiasm for courting females; not a good sign.
Increasing the generation interval allows for the removal of genetically inferior individuals. For
example, one male, which I had planned to use for breeding, looked normal at 6 months. At 7 months,
however, he developed “fin rot” (Fig 7). The slow tail erosion suggested early senescence, poor
circulation and diminished immunity to bacteria—all genetic weaknesses. I no longer considered him
for breeding. Instead, I used his equally attractive brother
who kept his tail intact throughout his life. By simply
waiting a couple months before using the males for
breeding, I could select out a tendency towards “fin rot”
in my male guppies.
In early 2020, I had narrowed a batch down to one
single female who even when young was noticeably
bigger and more vigorous than her sisters. When she was
7 months, I started saving her progeny hoping that she
would make the 8-month cut-off. She did and then some
(Fig 8). Fortunately, I managed to get three big litters
from her, all sired by different males. She represents the
Fig 8 One Year Plus Survival
3rd generation of females that I have selected for
This big beauty was my first female that
longevity. This female was a major improvement on the survived past one year. She grew to an
fish that I started out with in 2017. impressive 4.5 cm body length. Big females
As to the males, they are currently a “work in like this can easily produce 60 or more fry.
progress.” Their longevity has increased from the 4-5
months that I started out with in 2017 with my BG strain,
but monitoring their reproductive success is more difficult than with the females. It requires mating
them to virgins and seeing whether the females actually reproduce.
Setting a minimum threshold for the generation interval is critical. Currently, mine is 8 months.
Breeders must live at least 8 months before I will keep their progeny. If they die beforehand, I discard
their progeny. This threshold is based on the current situation in my guppies. In the future, I may be
able to increase it.
9

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.

Recommendations for Increasing Longevity


For those who want to improve the longevity in their guppies, here are my recommendations:

❖ 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).
10

References

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8. Discussion (October 2018) about longevity on the Guppy Gene Collectors forum
https://www.facebook.com/groups/GuppyGeneCollectors/
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Diana Walstad is the author of Ecology of the Planted Aquarium (2013). For articles
and more information about her books, see: http://dianawalstad.com

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