Elevational and Temporal Trends in Salix-Feeding Beetles and Associated Insects Along Three Sierra Nevada, California Drainages

Elevational And Temporal Trends In Salix-feeding Beetles And Associated Insects Along Three Sierra Nevada, California Drainages

John Smiley1, Elizabeth Dahlhoff2, and Nathan Rank3

1University of California White Mountain Research Station 2Santa Clara University 3Sonoma State University John Smiley and Nathan Rank at Upper Tyee Lake in Bishop Creek drainage

“team beetle”

Work supported by: National Science Foundation and our respective institutions Elizabeth Dahlhoff Elevation gradients are asymmetrical:

• Most species will have an upper limit beyond which they cannot physically survive (usually cold stress) • For many taxa, biodiversity decreases as elevation increases • Many species have a lower limit along the gradient, below which natural enemies or heat-related stress prevents survival. • Between these limits is a zone in which a species can survive, reproduce and send out dispersers to colonize new habitats.

drought or heat stress, predators, pathogens and competitors 1 cold stress zone of positive fitness mortality rate 0 low elevation high Q: how do plant and animal species respond to changing climate along steep, montane elevation gradients?

A: look at a multi-species food web inhabiting such a gradient, and study it for a long period of time. Central- eastern Sierra Nevada University of California White Mountain Research Station Owens Valley Laboratories Rock Creek

Bishop Creek

Big Pine Creek We have studied a willow-insect food chain since 1981. Salix orestera Sierra Willow

For this presentation, we will examine: changes along •Insect Range Distribution since 1981 elevation •Foliage Air Temperatures and trends gradients in 3 •Beetle Numbers since 1998 drainages •Beetle Genetics and Physiology beetle: Chrysomela aeneicollis Coleoptera: Chrysomelidae wasp: Symmorphus cristatus Hymenoptera: Vespidae

fly larva: Parasyrphus melanderi Ants: Formica sp. Diptera: Syrphidae Hymenoptera: Formicidae 3750

3500

3250

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2750 Y 2500

2250

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1750

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1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year Y upper limit lower limit

C. aeneicollis elevational range: Long term record from Big Pine Creek Are there decadal trends?: yes! 3750

3500

3250

3000

2750 Y 2500

2250

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1750

1500

1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year Y upper limit lower limit

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175

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125

snow index 100

50 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year

C. aeneicollis elevational range and winter snow % of normal: Long term record from Big Pine Creek & Owens Valley region Behavioral ecology of wasp: Symmorphus cristatus at Falls site, 2900m, Big Pine Creek

Lodgepole pine borer (Family Carambycidae) 3750

3500

3250

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2750 Y 2500

2250

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1750

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1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year Y upper limit lower limit

S. cristatus elevational range: Long term record from Big Pine Creek (note: prey most abundant along upper orange band) Quantitative C. aeneicollis counts begun in 1998, Quantitative foliage air temperature recording begun in 2000.

3750

3500

3250

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2750 Y 2500

2250

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1750

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1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year Y upper limit lower limit White plastic cups were used as radiation shields, hung upside down from 1- 2 cm diameter willow branch, with HOBO temp. logger suspended inside.

Shown: S. orestera above Green Lake, elev. 3410m (11,200’) Big Pine drainage

Elevation color bands: 4000-4250m light gray 3750-4000m light blue 3500-3750m blue 3250-3500m blue-green 3000-3250m green 2750-3000m yellow-green 2500-2750m tan 2250-2500m light tan 2000-2250m yellow Bishop Creek drainage

Elevation color bands: 4000-4250m light gray 3750-4000m light blue 3500-3750m blue 3250-3500m blue-green 3000-3250m green 2750-3000m yellow-green 2500-2750m tan 2250-2500m light tan 2000-2250m yellow

Daniel Pritchett 30

10 Y

-10

-20 01/01/2005 01/01/2006 01/01/2007 01/01/2000 01/01/2001 01/01/2002 01/01/2003 01/01/2004 date Y Mean(maxtemp) Mean(mintemp) Mean(avetemp)

Foliage air temperatures, averaged over 18 sites (= daily “weather”). Note snow burial periods in spring. Q: Do foliage air temperatures lapse with increasing elevation? What is the lapse rate?

A: The answer depends on the measurement (max or min) and the creek drainage. Foliage air temperature, all sites, all months:

Y 5

-5

-10

2600 2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp dev+13 mintemp dev -2 avetemp dev + 3.74 daily maximum lapse rate = 8.3 °C/km elev. daily average lapse rate = 2.6 °C/km elev. daily minimum lapse rate = n.s. Foliage air temperature, all sites, summer only:

Y 15

2600 2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp+25.5 mintemp+3.5 avetemp+12.3

daily maximum lapse rate = 5.8 °C/km elev. daily average lapse rate = 2.8 °C/km elev. daily minimum lapse rate = n.s. Big Pine Creek foliage air temperatures, summer only

Y 15

2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp+25.5 mintemp+3.5 avetemp+12.3

daily maximum lapse rate = (12°C/km elev.) daily average lapse rate = (4.6°C/km) daily minimum lapse rate n.s. Bishop Creek foliage air temperatures, summer only

Y 15

2700 2800 2900 3000 3100 3200 3300 Mean(m elev) Y maxtemp+25.5 mintemp+3.5 avetemp+12.3

daily maximum lapse rate = n.s. daily average lapse rate n.s. daily minimum lapse rate n.s. Rock Creek foliage air temperatures, summer only

Y 15

2600 2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp+25.5 mintemp+3.5 avetemp+12.3

daily maximum lapse rate n.s. daily average lapse rate = 2.1 °C/km daily minimum lapse rate n.s. Q: are there long term trends in foliage air temperatures?

A: maybe not in our data, nor in a longer data set from Big Pine Creek. Yet warming trends are seen in other local data sets…. Barcroft Field Station July Average (Daily Max, Min Air Temperatures)

50 Y 45

1950 1960 1970 1980 1990 2000 2010 year Y July max July min

Bishop Airport July Average

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100

Y 80

1940 1950 1960 1970 1980 1990 2000 2010 year Y average max average min Barcroft Field Station July Average

50 Y 45

1950 1960 1970 1980 1990 2000 2010 year Y July max July min Big Pine Creek snow station, July Average

70 65 60 55 50 Y 45 40 35 30 25

50 55 60 65 70 75 80 85 90 95 100 105 110 July Y Column 3 Column 4 July Averages 65

50 Y 45 Barcroft 40

1950 1960 1970 1980 1990 2000 2010 year Y July max July min

70 65 60 55 50 Y Big Pine Creek snow station 45 40 35 30 25

50 55 60 65 70 75 80 85 90 95 100 105 110 July Y Column 3 Column 4

Y 15 Foliage – all sites mean 10

1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 year Y Mean(maxtemp) Mean(mintemp) 1984 2006

North Palisades Glacier Q: are there trends in number of beetles over time? Over elevation?

A: it depends on the drainage…and something C. aeneicollis numbers were estimated by abrupt counting as many beetles as possible during a 10 happened in minute period, at each of about 40 sites, 2006 including all of the temperature logger sites. elevation of peak beetle abundance

3400

3300

3200 ) m BP ( 3100 n o BC 3000 RC evati el 2900

2800

2700 1998 2000 2001 2002 2003 2004 2005 2006 late 2006 numbers of beetles at 6 sites in Big Pine Creek

6 e r 5 ce sco n 4 a d n u 3 ab e l 2 et e b 1

0 3126 3246 3385 2800 2926 2997 elevation (m)

1998 2000 2001 2002 2003 2004 2005 2006 elevation of peak beetle abundance

3400

3300

3200 ) m BP ( 3100 n o BC 3000 RC evati el 2900

2800

2700 1998 2000 2001 2002 2003 2004 2005 2006 late 2006 5

score 2

1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 year

Green Lake beetle abundance 1998-2006 30 30 30

20 20 20 Y Y Y 10 10 10

0 0 0

150 160 170 180 190 200 210 220 230 240 250 150 160 170 180 190 200 210 220 230 240 250 150 160 170 180 190 200 210 220 230 240 250 day day day

Plot for year=2000 Plot for year=2001 Plot for year=2002

30 30 30

20 20 20 Y Y Y 10 10 10

0 0 0

150 160 170 180 190 200 210 220 230 240 250 150 160 170 180 190 200 210 220 230 240 250 150 160 170 180 190 200 210 220 230 240 250 day day day

Plot for year=2003 Plot for year=2004 Plot for year=2005

20 Y 10 Nothing unusual about summer 2006? 0

150 160 170 180 190 200 210 220 230 240 250 Heavy rain/hail in early July. day

Plot for year=2006 Q: can genetics, physiology, and/or behavior help explain what might be happening?

A: look at PGI, heat shock proteins and running performance after different types of temperature stress. PGI

•Phosphoglucose Isomerase (PGI) is an important metabolic enzyme involving cellular energy supplies including glycolysis and glucogenesis. •PGI is often observed to limit aerobic performance. •Studies on a variety of ectothermic species including sea anemones, butterflies and beetles (our study) suggest that alleles of PGI exhibit a tradeoff between enzymatic efficiency and thermo-stability: the more active, efficient isozymes seem to be more sensitive to temperature extremes. •In our system the PGI-1 enzyme is the more efficient but less stable form when compared with PGI-4. Rock Creek; Mt. Dana; Oregon; Montana; Colorado north 1 ▼▼▲ ▲▼ North Lake ▲ ●▼ ▲ ◆ 0.9 ▼ ▼ ▲ ▲ George Lake ◆ ▼ ▲▼ ● 0.8 Bishop ● ◆ 0.7 ◆ Creek ● ◆◆◆◆ Green Lake ● ● 0.6 ◆ ● ● Chocolate ● ● ◆ FREQUENCY 0.5 ◆ Lakes south 0.4 PGI-1 0.3 0.2 Big Pine Creek; Taboose Creek

0.1

2500 2700 2900 3100 3300 ELEVATION PGI, HSP’s and running performance

•Temperature stress in ectotherms such as the willow leaf beetle induces heat shock proteins (HSP’s) which quickly build up in the tissues and protect enzymes such as PGI from degrading and losing efficiency. •The induction of HSP’s permit better enzyme performance after stressful conditions, and thus faster running speed. •Our findings suggest that PGI-1 individuals seem to be more fit during milder weather with infrequent temperature extremes, while PGI-4 individuals are better in more extreme environments. •Cold temperature extremes are much more common in the early and late season, but the timing of snowmelt determines whether or not beetles experience this severity of cold. #Days logger is buried under snow, as function of elevation

200

150 s day 100

0 2800 2900 3000 3100 3200 3300 elev

X=Rock Creek; O=Big Pine Creek; =Bishop Creek Findings:

Foliage air temperatures: • Lapse with elevation as predicted for afternoon highs. • Lapse rate reduced for nighttime lows. Nighttime lapse rate “anomaly” seen most strongly in north-south oriented Rock Creek and Bishop Creek drainages • Minimal warming seen over past 10-20 years

Yet: Beetle distribution change: • 25 year inter-decadal upward trend in Big Pine Creek for beetles and predators. • 8-year upward trend in Big Pine Creek but not Bishop Creek or Rock Creek. Temperature regime? Wasp predation? • Upward shift in summer 2006 – will it persist? • Genetics and physiology: no clear pattern with elevation, but with latitude, yes. Questions

Why are beetles moving up? Why shift in 2006?

Temperature shifts not recorded/analyzed in data? Timing of snow melt? Earlier snow melt probably favors higher elevation beetles where the snow melts later.

Changing plant quality as food for beetles? Does altering ambient CO2 concentration affect Salix orestera foliage? Published literature would suggest not, but we should measure this if possible.

Need longer-term data set. We are trying to secure long-term funding from NSF. North Palisade Glacier: how long before the willows get here? Present glaciers END SLIDE 2001 Falls site, Big Pine Creek

2006

Lodgepole pine borer (Family Carambycidae)

Big Pine drainage

Elevation color bands: 4000-4250m light gray 3750-4000m light blue 3500-3750m blue 3250-3500m blue-green 3000-3250m green 2750-3000m yellow-green 2500-2750m tan 2250-2500m light tan 2000-2250m yellow

Daniel Pritchett

Big Pine Creek foliage air temperatures, all months

10 Y 5

-5

2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp dev+13 mintemp dev -2 avetemp dev + 3.74

daily maximum lapse rate = (9°C/km elev.) daily average lapse rate = (3°C/km) daily minimum lapse rate n.s. Bishop Creek foliage air temperatures all months

10 Y

-5

2700 2800 2900 3000 3100 3200 3300 Mean(m elev) Y maxtemp dev+13 mintemp dev -2 avetemp dev + 3.74

daily maximum lapse rate = (11°C/km) daily average lapse rate =(4°C/km) daily minimum lapse rate n.s. Rock Creek foliage air temperatures, all months

5 Y

-5

-10

2600 2700 2800 2900 3000 3100 3200 3300 3400 Mean(m elev) Y maxtemp dev+13 mintemp dev -2 avetemp dev + 3.74

daily maximum lapse rate = (5.5°C/km) daily average lapse rate n.s. daily minimum lapse rate n.s. (+4°C/km!) Barcroft Field Station July Max, Min Air Temperatures

50 Y 45

1950 1960 1970 1980 1990 2000 2010 year Y July max July min

Bishop Airport

110

100

Y 80

1940 1950 1960 1970 1980 1990 2000 2010 year Y average max average min Barcroft Field Station July Max, Min Air Temperatures

50 Y 45

1950 1960 1970 1980 1990 2000 2010 year Y July max July min Falls Site, Big Pine Creek July Max, Min Air Temperatures

70 65 60 55 50 Y 45 40 35 30 25

50 55 60 65 70 75 80 85 90 95 100 105 110 July Y Column 3 Column 4 North Palisade Glacier: how long before the willows get here? Present glaciers PGI, HSP’s and running performance

Temperature stress in ectotherms induces heat shock proteins (HSP’s) which quickly build up in the tissues and protect enzymes such as PGI from degrading and losing efficiency.

Our findings suggest that PGI-1 , the more sensitive form, immediately induces HSP production in response to only 4h of extreme heat (36 C.) or cold (-4 C.), but that this is not enough stimulus to induce drastic HSP production in beetles with PGI-4. As a result, PGI-1 beetles outperform (outrun) PGI-4 beetles after either stimulus. However, after a second bout of heat or cold on the second day, the PGI-4 beetles induce HSP’s and generally outperform the PGI-1 beetles thereafter. Thus PGI-1 seems to be superior during mild weather with infrequent temperature extremes, while PGI-4 is best in more extreme environments. Genetics

•Beetles are collected in the field, frozen and analyzed in the lab •Principal alleles (forms) of the enzyme are determined, including PGI-1 and PGI-4 •Predict that PGI-1 would be more common at lower elevations?