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Date: Thu, 1 Oct 2020 21:02:28 +0000 (UTC)
From: Kenneth Davenport via groups.io <kdavenport93306=<yahoo.com...>
Subject: Re: [NorWestLeps] San Francisco butterflies breeding times

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Content-Transfer-Encoding: quoted-printable

Best Wishes, Ken Davenport <kdavenport93306...> or flutterflies93306@=
att.net=20
=20
And another thing to consider in harmony with Paul's comments: John Emmel=
's old statement: Under a given set of circumstances, butterflies will do w=
hatever they darn well please.

I keep records of early and late flights for California, to my knowledge =
no one has ever kept breeding season records. Doing so would probably break=
California's extensive privacy laws.=20
On Thursday, October 1, 2020, 12:10:12 AM PDT, Paul Johnson via group=
s.io <pjpolliwog=<3Dyahoo.com...> wrote:
=20
I don't know enough about the butterflies in your area (or mine) to answe=
r your Vanessa question definitively.=C2=A0 Maybe nobody does.=C2=A0=20

Here's some food for thought, based somewhat on speculation...

When we speak about animal behavior, we are often speaking in generalities=
.=C2=A0 It's not that all individuals of a species act a certain way.=C2=A0=
It's more of a probability that an individual will act a certain way.=C2=
=A0=20

So, for example, we can say that the Monarchs that return to California in=
the fall have entered reproductive diapause, and will not start breeding u=
ntil late winter or early spring.=C2=A0 But every once in a while, we'll se=
e a mating pair of Monarchs in October.=C2=A0 So maybe 99% of Monarchs foll=
ow the pattern.=C2=A0 What determines that 99% number?=C2=A0 Is it affected=
by environmental factors such as weather or larval food conditions?=C2=A0 =
Or is in inborn, genetically based?=C2=A0 Even if it's inborn, it's possibl=
e that natural selection can change it over the years, so that if environme=
ntal changes favor breeding in the fall, over the years we may see more and=
more Monarchs breeding then.=C2=A0 So even if we "know" something about wh=
at a species does in SF this year, that might have little bearing on what t=
hey do in 10 years or 50 years.=C2=A0 And this ratio might also vary geogra=
phically.

We need to remember that butterflies probably can't predict the future, an=
d they're probably not adopting a particular strategy because they think it=
's going to work.=C2=A0 More likely, individual butterflies are predetermin=
ed to behave a certain way, and there is some variation between individuals=
.=C2=A0 Often what we probably see is that the strategy followed by most in=
dividuals in an area is the one that works most of the time, and strategies=
followed by fewer individuals fail most of the time, but in some years the=
se "failure" strategy followers are successful when the majority's strategy=
fails.=C2=A0 The butterflies have no idea whether their strategy will work=
.=C2=A0 They just live their lives the way they are programmed to, and some=
survive while others do not.=C2=A0 Those that survive pass their genes alo=
ng to the next generation.

Here's a completely made up example...

There is a butterfly called the California Lady that ranges from the Bay A=
rea to San Diego.=C2=A0 Its native host plants are gumweeds.=C2=A0 Now it t=
urns out that gumweeds can survive the winter in San Diego, but in the SF B=
ay Area the winters are so cold that gumweeds only survive as seeds.=C2=A0 =
Throughout their range, the butterflies can pass the winter as a caterpilla=
r or an adult, but the caterpillars only make it if they have plenty of fre=
sh gumweed to eat.=C2=A0 And overwintering adults are highly prone to attac=
k by ants as they hide out in cracks and crevices to shelter against cold w=
inter nights.=C2=A0=20

Now a detailed life history study from 1990-2000 found that in San Diego, =
90% of individuals overwinter as caterpillars and 10% overwinter as adults.=
=
=C2=A0 Once every decade, a very cold winter kills off the gumweed and the=
overwintering caterpillars all starve to death, while some of the overwint=
ering adults survive, despite many of them being eaten by ants.=C2=A0 Even =
though overwintering as a caterpillar is the successful strategy in Sand Di=
ego in most years, in very cold years the strategy of overwintering as an a=
dult is what keeps the species from being wiped out locally.=C2=A0 So it pa=
ys to have some outliers in the population, even though most years they fai=
l.

The same 1990-2000 study found that SF Bay Area California Lady butterflie=
s have the opposite strategy.=C2=A0 90% of individuals overwinter as adults=
and 10% as caterpillars.=C2=A0 In 9 out of 10 winters, the cold weather ki=
lls off the gumweed and all the overwintering caterpillars starve to death.=
=
=C2=A0 Only the overwintering adults survive the winter.=C2=A0 But once pe=
r decade, an unusually warm winter allows the gumweed to live through the w=
inter, and the overwintering caterpillars do much better than the overwinte=
ring adults, which get devoured by the ants, which are more active than usu=
al on the warmer nights.=C2=A0 So here, overwintering as an adult is usuall=
y the winning strategy, but in very warm years it's the overwintering cater=
pillars that keep the species from being wiped out locally.=C2=A0 Again, it=
pays to have some outliers in the population, even though they fail in mos=
t years.

The study was repeated in 2010-2020, but something had changed: Guernsey G=
umweed was introduced to California and although it did not do well in the =
face of competition from natives gumweeds in natural areas, it thrived in c=
ities.=C2=A0 The thing about Guernsey Gumweed is that it does quite well in=
Bay Area winters.=C2=A0 So what did the study find?=C2=A0 Things hadn't ch=
anged in San Diego: overwintering caterpillars were still the most successf=
ul strategy in most years.=C2=A0 And in natural areas in the East Bay Hills=
, they found the same results as in the previous study: overwintering adult=
s were the most successful in most years.=C2=A0 But in SF where Guernsey Gu=
mweed was well-established and supported caterpillars through most winters,=
overwintering California Lady caterpillars became the dominant strategy, j=
ust like in San Diego.=C2=A0=20

The study was then put on hold indefinitely because, after all, it was 202=
0.=C2=A0 But then in 2100-2110 it was repeated.=C2=A0 By then SF Bay area t=
emperatures had warmed by a few degrees, allowing native gumweeds to surviv=
e the winter in all but the coldest years.=C2=A0 And in San Diego, winter t=
emperatures were so warm that gumweeds never died off, not even in the cold=
est winters.=C2=A0 So what did the study find?=C2=A0 In San Diego, the over=
wintering adults had all but disappeared.=C2=A0 Every individual found in w=
inter was a caterpillar.=C2=A0 The same was true in SF, where the introduce=
d Guernsey Gumweed supported overwintering caterpillars even through the co=
ldest winters, and no overwintering adults could be found.=C2=A0 What about=
in the natural areas in the East Bay Hills, where the native gumweeds made=
it through 9 out of 10 winters?=C2=A0 There, the species had adopted the f=
ormer San Diego scenario, in which 90% of California Ladies overwintered as=
caterpillars and only 10% as adults.=C2=A0=20

Now, imagine if someone asked the question: Do California Ladies overwinte=
r as caterpillars or adults?=C2=A0 The answer would depend not only on the =
location, but also the year.=C2=A0 Even back in the 1990s, the answer depen=
ded on the year, with most years favoring overwintering caterpillars, but 1=
out of 10 years favoring overwintering adults.=C2=A0 Also, if someone had =
seen a California Lady in San Diego in June of 1995 and asked whether its o=
ffspring would overwinter as adults or caterpillars, you wouldn't have been=
able to answer definitively, although you could have said that there's a 9=
0% chance they will overwinter as caterpillars and a 10% chance they'll ove=
rwinter as adults.=C2=A0=20

Now back to the real world: Do American Ladies in SF breed in September?=
=C2=A0 How many butterflies would we need to observe to answer that questi=
on, and over how many years?=C2=A0 And if we had studied them in the East B=
ay Hills, could we apply our results to the same species living in SF?=C2=
=A0 Or if we had studied them before the arrival of the non-native cudweed=
, would our results still apply after the cudweed had been established for =
a while?=C2=A0 Will our results still apply as average temperatures change?

Forgetting all of that for a moment, there are other reasons why answering=
the "breeding season(s)" question may be difficult...

For butterflies with multiple generations per year (all Vanessas in SF?), =
the timing of those generations may vary each year, and they may not always=
be in sync.=C2=A0 (The following example is made up.)=C2=A0 Maybe one Vane=
ssa species usually breeds in April (1st generation), June (2nd generation)=
, and August (3rd generation), and then goes into reproductive diapause unt=
il April.=C2=A0 But in an especially warm year, maybe that's March, May, an=
d July.=C2=A0 And maybe some of those July butterflies do breed and have a =
generation that emerges in September and then diapauses through the winter.=
=
=C2=A0 And maybe in a cold year they start in May and only get two generat=
ions in that year.=C2=A0=20

Some butterflies have partial generations.=C2=A0 For example, Sara Orange-=
tips apparently have a full generation that flies in early spring, and some=
of their offspring emerge in late spring while others wait until the follo=
wing spring to emerge.=C2=A0 So their "breeding periods" may be easy to ide=
ntify, but only some individuals are breeding during the second breeding pe=
riod.

Butterflies may have their life histories in sync with the life history of=
their local host plant, which may vary over small geographic scales.=C2=A0=
For example, Anise Swallowtails living in towns have been shown to have mo=
dified the timing of their life history to match the timing of non-native F=
ennel, while Anise Swallowtails in nearby natural areas retain their natura=
l phenology.=C2=A0 So the breeding seasons of Anise Swallowtails can be dif=
ferent in areas just a few miles apart.=C2=A0=20

And as another example, I suspect that the Variable Checkerspots in my are=
a have such a long flight season because their caterpillars feed on at leas=
t three different plants, each of which grows in a different microclimate a=
nd has a different phenology.=C2=A0 So when defining the breeding season fo=
r this species, should we separate out each host plant cohort, or should we=
just lump them all together?=C2=A0=20

I don't have any real data on this, but I have wondered about the Red Admi=
rals I can find hilltopping on just about any warm winter day.=C2=A0 The pu=
rpose of hilltopping is mating.=C2=A0 Males go to hilltops to wait for rece=
ptive females to arrive, and then they mate.=C2=A0 So if I see male Red Adm=
irals on hilltops, it must be their breeding season, right?=C2=A0 Maybe not=
.=C2=A0 Perhaps males are just covering their bases: any day that's warm en=
ough to fly, they'll head for a nearby hilltop just in case a receptive fem=
ale might arrive.=C2=A0 But maybe females won't emerge and head for hilltop=
s until March.=C2=A0 If that's the case, then even though males appear to b=
e ready to breed all winter long, I don't think it's really breeding season=
until receptive females join them.=C2=A0 But let's remember that there are=
often some outliers that don't follow the "normal" pattern.=C2=A0 What if =
1% of females are receptive throughout the winter and will fly to hilltops =
on warm winter days to mate?=C2=A0 Does that 1% make winter into Red Admira=
l breeding season?=C2=A0 What if 10% of female Red Admirals breed in winter=
?=C2=A0 50%?=C2=A0 How do we determine what percentage of butterflies in br=
eeding status is required to call it breeding season?

These examples are meant to convey some of the complexities of determining=
the breeding season(s) of butterfly species.=C2=A0 Every species is differ=
ent, and there may be complications, variability, local variation, evolutio=
n, etc.=C2=A0 And definitions of "breeding season" may not be as straightfo=
rward as we would hope.

I hope this helps!=C2=A0=20

I encourage folks to photograph butterflies nectaring, breeding, laying eg=
gs, etc. and post their observations on iNaturalist.=C2=A0 The more of thes=
e observations get posted, the more data will be available to investigate t=
hese kinds of questions.=C2=A0=20

Thanks,

Paul
--=20
Paul G. Johnson
Paicines, CA
<pjpolliwog...>


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------=_Part_960525_2045919659.1601586148218
Content-Type: text/html; charset=UTF-8
Content-Transfer-Encoding: quoted-printable

<div> Best Wishes, Ken Davenport <kdavenport93306...> o=
r <flutterflies93306...> <br> <br> And another thing to consi=
der in harmony with Paul&#39;s comments: John Emmel&#39;s old statement: Un=
der a given set of circumstances, butterflies will do whatever they darn we=
ll please.<br><br> I keep records of early and late flights for Californi=
a, to my knowledge no one has ever kept breeding season records. Doing so w=
ould probably break California&#39;s extensive privacy laws. <br> =
</div> <div class=3D"yahoo_quoted" style=3D"margin:1=
0px 0px 0px 0.8ex;border-left:1px solid #ccc;padding-left:1ex;"> =
<div style=3D"font-family:'Helvetica Neue', Helvetica, Arial, =
sans-serif;font-size:13px;color:#26282a;"> <=
div> On Thursday, October 1, 2020, 12:10:12 AM PDT, Paul=
Johnson via groups.io &lt;pjpolliwog=<3Dyahoo.com...>&gt; wrote: =
</div> <div><br></div> <div><br></=
div> <div><div id=3D"yiv2487590064">I don't know enough abou=
t the butterflies in your area (or mine) to answer your Vanessa question de=
finitively.&nbsp; Maybe nobody does.&nbsp; <br><br>Here's some food for tho=
ught, based somewhat on speculation...<br><br>When we speak about animal be=
havior, we are often speaking in generalities.&nbsp; It's not that all indi=
viduals of a species act a certain way.&nbsp; It's more of a probability th=
at an individual will act a certain way.&nbsp; <br><br>So, for example, we =
can say that the Monarchs that return to California in the fall have entere=
d reproductive diapause, and will not start breeding until late winter or e=
arly spring.&nbsp; But every once in a while, we'll see a mating pair of Mo=
narchs in October.&nbsp; So maybe 99% of Monarchs follow the pattern.&nbsp;=
What determines that 99% number?&nbsp; Is it affected by environmental fac=
tors such as weather or larval food conditions?&nbsp; Or is in inborn, gene=
tically based?&nbsp; Even if it's inborn, it's possible that natural select=
ion can change it over the years, so that if environmental changes favor br=
eeding in the fall, over the years we may see more and more Monarchs breedi=
ng then.&nbsp; So even if we "know" something about what a species does in =
SF this year, that might have little bearing on what they do in 10 years or=
50 years.&nbsp; And this ratio might also vary geographically.<br><br>We n=
eed to remember that butterflies probably can't predict the future, and the=
y're probably not adopting a particular strategy because they think it's go=
ing to work.&nbsp; More likely, individual butterflies are predetermined to=
behave a certain way, and there is some variation between individuals.&nbs=
p; Often what we probably see is that the strategy followed by most individ=
uals in an area is the one that works most of the time, and strategies foll=
owed by fewer individuals fail most of the time, but in some years these "f=
ailure" strategy followers are successful when the majority's strategy fail=
s.&nbsp; The butterflies have no idea whether their strategy will work.&nbs=
p; They just live their lives the way they are programmed to, and some surv=
ive while others do not.&nbsp; Those that survive pass their genes along to=
the next generation.<br><br>Here's a completely made up example...<br><br>=
There is a butterfly called the California Lady that ranges from the Bay Ar=
ea to San Diego.&nbsp; Its native host plants are gumweeds.&nbsp; Now it tu=
rns out that gumweeds can survive the winter in San Diego, but in the SF Ba=
y Area the winters are so cold that gumweeds only survive as seeds.&nbsp; T=
hroughout their range, the butterflies can pass the winter as a caterpillar=
or an adult, but the caterpillars only make it if they have plenty of fres=
h gumweed to eat.&nbsp; And overwintering adults are highly prone to attack=
by ants as they hide out in cracks and crevices to shelter against cold wi=
nter nights.&nbsp; <br><br>Now a detailed life history study from 1990-2000=
found that in San Diego, 90% of individuals overwinter as caterpillars and=
10% overwinter as adults.&nbsp; Once every decade, a very cold winter kill=
s off the gumweed and the overwintering caterpillars all starve to death, w=
hile some of the overwintering adults survive, despite many of them being e=
aten by ants.&nbsp; Even though overwintering as a caterpillar is the succe=
ssful strategy in Sand Diego in most years, in very cold years the strategy=
of overwintering as an adult is what keeps the species from being wiped ou=
t locally.&nbsp; So it pays to have some outliers in the population, even t=
hough most years they fail.<br><br>The same 1990-2000 study found that SF B=
ay Area California Lady butterflies have the opposite strategy.&nbsp; 90% o=
f individuals overwinter as adults and 10% as caterpillars.&nbsp; In 9 out =
of 10 winters, the cold weather kills off the gumweed and all the overwinte=
ring caterpillars starve to death.&nbsp; Only the overwintering adults surv=
ive the winter.&nbsp; But once per decade, an unusually warm winter allows =
the gumweed to live through the winter, and the overwintering caterpillars =
do much better than the overwintering adults, which get devoured by the ant=
s, which are more active than usual on the warmer nights.&nbsp; So here, ov=
erwintering as an adult is usually the winning strategy, but in very warm y=
ears it's the overwintering caterpillars that keep the species from being w=
iped out locally.&nbsp; Again, it pays to have some outliers in the populat=
ion, even though they fail in most years.<br><br>The study was repeated in =
2010-2020, but something had changed: Guernsey Gumweed was introduced to Ca=
lifornia and although it did not do well in the face of competition from na=
tives gumweeds in natural areas, it thrived in cities.&nbsp; The thing abou=
t Guernsey Gumweed is that it does quite well in Bay Area winters.&nbsp; So=
what did the study find?&nbsp; Things hadn't changed in San Diego: overwin=
tering caterpillars were still the most successful strategy in most years.&=
nbsp; And in natural areas in the East Bay Hills, they found the same resul=
ts as in the previous study: overwintering adults were the most successful =
in most years.&nbsp; But in SF where Guernsey Gumweed was well-established =
and supported caterpillars through most winters, overwintering California L=
ady caterpillars became the dominant strategy, just like in San Diego.&nbsp=
; <br><br>The study was then put on hold indefinitely because, after all, i=
t was 2020.&nbsp; But then in 2100-2110 it was repeated.&nbsp; By then SF B=
ay area temperatures had warmed by a few degrees, allowing native gumweeds =
to survive the winter in all but the coldest years.&nbsp; And in San Diego,=
winter temperatures were so warm that gumweeds never died off, not even in=
the coldest winters.&nbsp; So what did the study find?&nbsp; In San Diego,=
the overwintering adults had all but disappeared.&nbsp; Every individual f=
ound in winter was a caterpillar.&nbsp; The same was true in SF, where the =
introduced Guernsey Gumweed supported overwintering caterpillars even throu=
gh the coldest winters, and no overwintering adults could be found.&nbsp; W=
hat about in the natural areas in the East Bay Hills, where the native gumw=
eeds made it through 9 out of 10 winters?&nbsp; There, the species had adop=
ted the former San Diego scenario, in which 90% of California Ladies overwi=
ntered as caterpillars and only 10% as adults.&nbsp; <br><br>Now, imagine i=
f someone asked the question: Do California Ladies overwinter as caterpilla=
rs or adults?&nbsp; The answer would depend not only on the location, but a=
lso the year.&nbsp; Even back in the 1990s, the answer depended on the year=
, with most years favoring overwintering caterpillars, but 1 out of 10 year=
s favoring overwintering adults.&nbsp; Also, if someone had seen a Californ=
ia Lady in San Diego in June of 1995 and asked whether its offspring would =
overwinter as adults or caterpillars, you wouldn't have been able to answer=
definitively, although you could have said that there's a 90% chance they =
will overwinter as caterpillars and a 10% chance they'll overwinter as adul=
ts.&nbsp; <br><br>Now back to the real world: Do American Ladies in SF bree=
d in September?&nbsp; How many butterflies would we need to observe to answ=
er that question, and over how many years?&nbsp; And if we had studied them=
in the East Bay Hills, could we apply our results to the same species livi=
ng in SF?&nbsp; Or if we had studied them before the arrival of the non-nat=
ive cudweed, would our results still apply after the cudweed had been estab=
lished for a while?&nbsp; Will our results still apply as average temperatu=
res change?<br><br>Forgetting all of that for a moment, there are other rea=
sons why answering the "breeding season(s)" question may be difficult...<br=
><br>For butterflies with multiple generations per year (all Vanessas in SF=
?), the timing of those generations may vary each year, and they may not al=
ways be in sync.&nbsp; (The following example is made up.)&nbsp; Maybe one =
Vanessa species usually breeds in April (1st generation), June (2nd generat=
ion), and August (3rd generation), and then goes into reproductive diapause=
until April.&nbsp; But in an especially warm year, maybe that's March, May=
, and July.&nbsp; And maybe some of those July butterflies do breed and hav=
e a generation that emerges in September and then diapauses through the win=
ter.&nbsp; And maybe in a cold year they start in May and only get two gene=
rations in that year.&nbsp; <br><br>Some butterflies have partial generatio=
ns.&nbsp; For example, Sara Orange-tips apparently have a full generation t=
hat flies in early spring, and some of their offspring emerge in late sprin=
g while others wait until the following spring to emerge.&nbsp; So their "b=
reeding periods" may be easy to identify, but only some individuals are bre=
eding during the second breeding period.<br><br>Butterflies may have their =
life histories in sync with the life history of their local host plant, whi=
ch may vary over small geographic scales.&nbsp; For example, Anise Swallowt=
ails living in towns have been shown to have modified the timing of their l=
ife history to match the timing of non-native Fennel, while Anise Swallowta=
ils in nearby natural areas retain their natural phenology.&nbsp; So the br=
eeding seasons of Anise Swallowtails can be different in areas just a few m=
iles apart.&nbsp; <br><br>And as another example, I suspect that the Variab=
le Checkerspots in my area have such a long flight season because their cat=
erpillars feed on at least three different plants, each of which grows in a=
different microclimate and has a different phenology.&nbsp; So when defini=
ng the breeding season for this species, should we separate out each host p=
lant cohort, or should we just lump them all together?&nbsp; <br><br>I don'=
t have any real data on this, but I have wondered about the Red Admirals I =
can find hilltopping on just about any warm winter day.&nbsp; The purpose o=
f hilltopping is mating.&nbsp; Males go to hilltops to wait for receptive f=
emales to arrive, and then they mate.&nbsp; So if I see male Red Admirals o=
n hilltops, it must be their breeding season, right?&nbsp; Maybe not.&nbsp;=
Perhaps males are just covering their bases: any day that's warm enough to=
fly, they'll head for a nearby hilltop just in case a receptive female mig=
ht arrive.&nbsp; But maybe females won't emerge and head for hilltops until=
March.&nbsp; If that's the case, then even though males appear to be ready=
to breed all winter long, I don't think it's really breeding season until =
receptive females join them.&nbsp; But let's remember that there are often =
some outliers that don't follow the "normal" pattern.&nbsp; What if 1% of f=
emales are receptive throughout the winter and will fly to hilltops on warm=
winter days to mate?&nbsp; Does that 1% make winter into Red Admiral breed=
ing season?&nbsp; What if 10% of female Red Admirals breed in winter?&nbsp;=
50%?&nbsp; How do we determine what percentage of butterflies in breeding =
status is required to call it breeding season?<br><br>These examples are me=
ant to convey some of the complexities of determining the breeding season(s=
) of butterfly species.&nbsp; Every species is different, and there may be =
complications, variability, local variation, evolution, etc.&nbsp; And defi=
nitions of "breeding season" may not be as straightforward as we would hope=
.<br><br>I hope this helps!&nbsp; <br><br>I encourage folks to photograph b=
utterflies nectaring, breeding, laying eggs, etc. and post their observatio=
ns on iNaturalist.&nbsp; The more of these observations get posted, the mor=
e data will be available to investigate these kinds of questions.&nbsp; <br=
><br>Thanks,<br><br>Paul<br>-- <br>Paul G. Johnson<br>Paicines, CA<br><a hr=
ef=3D"mailto:pjpolliwog@yahoo," target=3D"_blank" rel=3D"nofollow" ymailto=
=3D"mailto:pjpolliwog@yahoo,"><pjpolliwog...></a> </div></div> =
</div> </div>


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