Anu Valtonen, Matthew P. Ayres, Heikki Roininen, Juha Pöyry & Reima Leinonen

Environmental controls on the phenology of moths ­ predicting plasticity and constraint under climate change

Anu Valtonen, Matthew P. Ayres, Heikki Roininen, Juha Pöyry & Reima Leinonen

Electronic Supplemental Material

Online Resource 1. For two sites the daily maximum and minimum temperatures were estimated using the nearest weather station records adjusted for temperature lapse rates as: T = b0+b1Lat+b2Alt, where b0 = temperature at the weather station, Lat and Alt = difference in latitude and elevation between the weather station and the trapping location, and b1 and b2 = lapse rates for latitude and elevation estimated from analyses of the 35 weather stations across the country. The average lapse rates (b1 and b2) were -0.492 °C / 100 km and -0.490 °C / 100 m for maximum temperatures and -0.664 °C / 100 km and -0.418 °C / 100 m for minimum temperatures. However, the lapse rates varied seasonally, being greatest in mid-winter and smallest in mid-summer (presumably due to lower mixing of air in the winter). We adjusted for this seasonality by making b1 and b2 sin functions of the solar year (annual minimum to maximum lapse rates were -0.389 to -0.633 °C / 100 km and -0.288 to -0.657 °C / 100 m for maximum daily temperatures and -0.337 to -1.104 °C / 100 km and -0.406 to -0.462 °C / 100 m for minimum daily temperatures). We also, initially, compared the lapse rates between maritime and continental sites and found differences in lapse rates for altitude. However, for simplicity we did not include different lapse rates for minimum and maximum temperatures for each month and for maritime and continental sites separately, because the (only) maritime site for which the temperatures had to be estimated differed only maximum of 14 meters in altitude from the weather stations used to estimate the temperatures.

Online Resource 2. We used a modified version of Cesaraccio et al.’s (2001) model designed to estimate the hourly temperatures based on daily minimum and maximum temperature data. Sunset and sunrise hours for each day at each site were accessed from Astronomical Applications Department of the U.S. Naval Observatory (http://aa.usno.navy.mil/data/docs/RS_OneYear.php). The model was parameterized to fit the northern conditions by using nine years (1997-2005) of hourly temperature data from Punkaharju Forest Research Station in Eastern Finland (N 61° 48’, E 29° 19’). The hour of the maximum temperature was estimated in the following way: if the sunset occurred before 3 pm, then maximum temperature was reached one hour before sunset, and if the sunset occurred after 3 pm, maximum temperature was reached at 2 pm (October-January) or 3 pm (February-September). The temperature at sunset (To) was calculated based on daily maximum temperature (Tx) and the minimum temperature of the following day (Tp) as To = Tx – c(Tx - Tp), where c was estimated empirically from the hourly temperature records at Punkaharju: 0.17 (Jan), 0.11 (Feb), 0.23 (Mar), 0.45 (Apr), 0.50 (May), 0.69 (Jun), 0.67 (Jul), 0.62 (Aug), 0.50 (Sep), 0.38 (Oct), 0.26 (Nov) and 0.21 (Dec). The model was further modified to estimate the temperatures in the mid-night sun conditions by using only two sine-functions, i.e., from the time when sun is at its lowest to the time when it is at its highest and lowest again.

Online Resource 3. Literature used for ecological classification of species.

Huldén L, Albrecht A, Itämies J, Malinen P, Wettenhovi J (2000) Atlas of Finnish Macrolepidoptera. Suomen Perhostutkijain seura, Luonnontieteellinen keskusmuseo, Helsinki

Marttila O, Saarinen K, Haahtela T, Pajari M (1996) Suomen kiitäjät ja kehrääjät. Kirjayhtymä, Helsinki

Mikkola K, Jalas I (1977) Yökköset 1. Otava, Keuruu

Mikkola K, Jalas I (1979) Yökköset 2. Otava, Keuruu

Mikkola K, Jalas I, Peltonen O (1985) Mittarit 1. Suomen Perhostutkijain seura, Tampere

Mikkola K, Jalas I, Peltonen O (1989) Mittarit 2. Recallmed, Hanko

Seppänen EJ (1954) Suomen perhostoukkien ravintokasvit. WSOY, Porvoo

Online Resource 4. Formulas and models used in data-analyses.

obs = observed day of peak flight

pred = model predicted day of peak flight

K = number of estimable parameters in an approximating model; 1 in Sday model (σ2), 2 in Photoperiod models (σ2, threshold), 3 in Tsum models (σ2, base, threshold), 4 in Tsum∩Photoperiod models (σ2, base, two thresholds), +2 when TotTsum added (intercept, slope), + 1 + df of Pops when populations added (intercept, slopes of populations -1).

p = number of predictors; in additive models number of parameters +1 for the intercept.

Models used to test for variation among sites, i.e., population differentiation:

obs = pred + a + b*TotTsum (or equally: obs - pred = a + b*TotTsum)

obs = pred + a + b*Pop (or equally: obs - pred = a + b*Pop)

TotTsum = average annual total thermal sum of the site, calculated from the entire study period with base = 5 °C, starting and ending at the winter solstice

Pop = site #, representing populations

a and b = estimated parameters in the models
Online Resource 5. Schematic of the model selection procedure.

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Online Resource 6. Comparisons of alternative models of phenological controls for each of 112 species, with the best model(s) in bold face. The competing models were based on thermal sum (T), photoperiod (P), thermal sum modified by photoperiod (TP), and solar day (S). Results are summarized in column labelled “Class”; models separated by commas were not distinguishable based on the information; evidence for population differentiation in thermal or photoperiodic controls indicated with “(diff)”. Family: Dre = Drepanidae, Geo = Geometridae, Noc = Noctuidae, Arc = Arctiidae, Hep = Hepialidae, Not = Notodontidae, Las = Lasiocampidae. Peak = earliest-average-latest peak date of flight across all sites and years (day.month.). OW = overwintering stage: FGL = full-grown larva, SL = small larva, L = larva, E = egg, P = pupa, I = imago, Sev = several. Diet: M = monophagous, SO = strongly oligophagous, O = oligophagous, P = polyphagous. Global = global distribution: P = palearctic, WP = west palearctic, H = holarctic, NA = information not available. Local = local distribution pattern in Finland: S = South Finland, SC = South – Central Finland, SCN = South – North Finland. N = number of site-year combinations, from which the peak flight was calculated. Area = geographical “zones” covered in this study: A = South Finland, B = South-Central Finland, C = North-Central and North Finland. SST = total variation in peak flight. Δi = AICc (i) – minAICc and RMSE were not calculated for photoperiod when information for that species was restricted to one area (because photoperiod was redundant with solar day). Notes: * all models not comparable either because the “best” critical day length of Photoperiod model was so high that it was not reached at some of the southern sites or because the apparent threshold thermal sum was so high that it was not reached at some of the northern sites at some years.

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Achlya flavicornis Dre 12.4.-22.4.-15.5. P O P SCN 28 3 12 AB 1577 Base -4, start 12 202.2 0.0 4.7 TP

Sday 123.0 20.2 7.6

Photoperiod 15.4 26.6 8.2

Base 3 43.5 27.4 8.0

Agriopis aurantiaria Geo 30.9.-18.10.-3.11. E P WP SC 21 3 10 A 1311 Sday 301.6 0.0 8.1 P,S

Photoperiod 9.9

Base -5, start 15 h 3238.7 38.3 16.5

Base -5 3753.5 38.6 17.9

Agrochola litura Noc 12.9.-18.9.-24.9. E P WP S 11 1 11 A 118 Sday 271.8 0.0 3.4 P,S

Photoperiod 12.5

Base -5, start 15 h 2798.6 33.5 9.0

Base -5 3360.2 33.7 11.6

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Agrotis clavis Noc 1.7.-10.7.-25.7. SL P P SCN 15 2 11 AB 754 Base 6 518.1 0.0 5.1 T

Base 6, start 15 h 509.3 3.3 5.0

Sday 201.9 4.8 7.3

Photoperiod 18.8 24.8 13.0

Alcis repandatus Geo 4.5.-15.7.-8.8. SL P WP SC 46 6 12 AB 8089 Sday 206.4 0.0 13.4 S,T

Base 4 720.2 1.7 13.0

Base -5, start 16 h 1377.0 2.9 12.8

Photoperiod 18.3 15.3 15.5

Allophyes oxyacanthae Noc 25.8.-6.9.-25.9. E P P SC 21 3 12 AB 2111 Sday 259.1 0.0 10.3 S,TP

Base -5, start 16 h 2486.3 1.7 8.7

Photoperiod 13.8 2.9 10.8

Base 2 1655.0 15.3 10.8

Amphipoea fucosa Noc 19.7.-12.8.-8.9. E O P SCN 85 9 12 ABC 6732 Base -5, start 16 h 2013.4 0.0 8.4 TP

Sday 235.0 5.0 9.0

Photoperiod 16.2 13.7 9.3

Base 0 1616.3 39.6 10.7

Amphipoea oculea Noc 5.8.-23.8.-15.9. E O P SC 47 5 12 AB 4051 Sday 245.2 0.0 9.4 S

Base -5, start 16 h 2253.9 9.7 9.7

Photoperiod 15.1 11.5 10.4

Base 1 1704.9 31.5 12.5

Apamea sublustris Noc 25.6.-10.7.-18.7. SL O WP S 11 1 11 A 470 Base 5 574.1 0.0 4.3 T

Sday 201.7 3.2 6.9

Photoperiod 18.3

Base 6, start 15 h 482.7 4.9 4.3

Aplocera praeformata Geo 14.7.-24.7.-8.8. SL SO WP SC 15 2 10 AB 936 Base 2 1063.4 0.0 4.9 T

Base 1, start 8 h 1170.9 3.0 4.8

Sday 215.1 9.4 8.2

Photoperiod 17.8 16.4 9.4

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Autographa jota Noc 7.7.-20.7.-1.8. SL P P S 10 1 10 A 625 Base -2 1598.5 0.0 4.9 T

Sday 211.9 3.3 8.3

Photoperiod 17.7

Base -5, start 6 h 2103.0 4.3 4.4

Brachylomia viminalis Noc 11.8.-23.8.-5.9. E SO P SCN 11 2 6 BC 802 Sday 245.0 0.0 9.0 S,P,T

Photoperiod 15.4 1.2 8.2

Base -1 1775.3 1.6 7.0

Base -4, start 12 2175.2 3.6 6.0

Cerapteryx graminis Noc 14.7.-30.7.-20.8. E O P SCN 103 10 12 ABC 5620 Base -5, start 17 + Pop 0.0 4.6 TP(diff)

Base -5, start 17 1603.7 24.3 5.9

Sday 221.2 65.9 7.4

Base 3 1022.2 89.4 8.2

Photoperiod 17.7 140.5 10.6

Cerastis rubricosa Noc 15.4.-4.5.-23.5. P P P SCN 43 5 12 AB 2967 Base -1 259.6 0.0 6.2 T

Base -2, start 8 290.8 2.3 6.2

Sday 134.4 21.2 8.4

Photoperiod 16.5 36.3 9.8

Chersotis cuprea Noc 28.7.-5.8.-15.8. SL P P SC 39 5 12 BC 746 Sday 227.6 0.0 4.4 P,S

Photoperiod 17.3 1.7 4.4

Base -5, start 16 1823.6 14.5 4.9

Base 0 1396.3 40.1 7.0

Chortodes fluxus Noc 21.7.-30.7.-15.8. SL SO P SC 20 2 12 A 994 Base -2 1777.1 0.0 5.5 T

Base -3, start 6 1941.0 2.9 5.4

Sday 221.0 5.9 7.2

Photoperiod 17.0

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Colostygia pectinataria Geo 24.6.-12.7.-25.7. FGL SO P SC 23 3 12 AB 1799 Base 5 626.0 0.0 6.1 T,TP

Base 3, start 15 h 760.9 1.5 5.9 a

Sday 203.3 13.4 9.0

Photoperiod 18.5 22.7 10.5

Cosmia trapezina Noc 4.8.-16.8.-29.8. E P WP SC 21 3 11 AB 834 Base -2 2060.5 0.0 5.4 T,P,TP

Photoperiod 15.6 0.8 5.8

Base -4, start 9 h 2326.3 1.5 5.2

Sday 238.8 2.5 6.5

Crocallis elinguaria Geo 21.7.-1.8.-22.8. Sev P P SC 73 8 12 ABC 3384 Base -2, start 16 h + Pop 0.0 4.0 TP(diff)

Base -2, start 16 h 1488.2 4.1 4.8

Base 2 1190.9 36.1 6.0

Sday 223.8 50.7 6.9

Photoperiod 17.1 71.5 7.8

Cybosia mesomella Arc 21.6.-5.7.-25.7. SL P P SC 19 2 12 AB 1156 Base 8 339.8 0.0 3.1 T,TP

Base 8, start 15 h 336.5 1.3 2.9

Sday 196.7 30.8 8.0

Photoperiod 19.0 60.1 16.2

Dasypolia templi Noc 3.9.-12.9.-26.9. I O WP SCN 19 2 12 BC 540 Sday 265.7 0.0 5.5 P,S

Photoperiod 13.3 2.2 5.4

Base -5, start 17 h 2409.9 21.1 7.6

Base -5 2800.9 27.9 9.9

Diarsia brunnea Noc 1.7.-20.7.-8.8. SL P P SC 17 3 10 AB 2286 Base 6 622.0 0.0 8.2 T

Base 6, start 14 620.7 3.1 8.1

Sday 211.3 7.1 12.0

Photoperiod 18.0 14.2 13.7

a: base 5, start 15h and base 3, start 15h give the same fit

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Diarsia dahlii Noc 30.7.-13.8.-30.8. SL P P SC 35 4 12 B 1344 Sday 235.6 0.0 8.2 P,S

Photoperiod 16.2

Base -5, start 15 h 2148.2 20.4 12.0

Base -5 2442.9 26.2 13.7

Diarsia mendica Noc 25.6.-11.7.-12.8. SL P P SCN 52 6 12 ABC 6275 Sday 202.1 0.0 11.1 S,TP,T

Base -5, start 13 h 1507.0 1.1 10.5

Base 0 1042.1 1.9 10.8

Photoperiod 19.0 21.3 13.3

Dichonia aprilina Noc 8.9.-17.9.-25.9. E P P S 11 1 11 A 255 Sday 270.9 0.0 5.0 P,S

Photoperiod 12.6

Base -5, start 12 h 2980.9 22.3 8.0

Base -3 2821.1 22.4 10.2

Dysstroma citratum Geo 2.8.-20.8.-8.9. E SO H SCN 153 13 12 ABC 8361 Sday 243.0 0.0 7.4 S

Base -5, start 18 h 1914.0 40.0 8.3

Photoperiod 15.6 52.9 8.8

Base -5 2551.2 260.2 17.1

Dysstroma latefasciatum Geo 25.7.-4.8.-15.8. SL P P SC 21 3 11 BC 477 Sday 226.5 0.0 4.9 S

Photoperiod 17.4 5.9 5.3

Base -5, start 15 h 1881.7 12.2 5.4

Base 3 1038.2 17.8 6.6

Eilema complanum Arc 22.7.-3.8.-18.8. SL P P SC 27 3 12 A 1493 Base -1 1697.0 0.0 6.2 T,TP

Base -5, start 6 h 2417.4 1.6 6.0

Sday 225.3 6.4 7.6

Photoperiod 16.6

Eilema lurideolum Arc 14.7.-25.7.-15.8. SL P P SC 29 3 12 A 1739 Base 0 1382.0 0.0 4.7 T

Base 4, start 16 h 823.9 2.1 4.7

Sday 216.9 25.2 7.9

Photoperiod 17.3

Species Family Peak OW Diet Global Local N Sites Years Area SST Models ranked Threshold Δi RMSE Class Notes

Eilema lutarellum Arc 15.7.-24.7.-8.8. SL P P SC 23 3 12 AB 970 Base 1, start 15 h 1114.6 0.0 3.5 T,TP

Base 1 1200.0 1.0 3.8 b

Sday 215.9 21.1 6.6

Photoperiod 17.6 32.0 8.0

Enargia paleacea Noc 5.8.-21.8.-8.9. E P P SCN 32 6 9 ABC 1908 Sday 243.2 0.0 7.8 S,P,TP

Photoperiod 15.4 3.8 8.0

Base -3, start 16 h 1891.4 6.5 7.7