Daniel A. Herms
This report presents the phenological sequence of 56 plant and 22 insect species for Wooster, Ohio, in 1997. A phenological sequence developed in Midland, Michigan, based on five years of data was evaluated for its accuracy in predicting insect activity in Wooster in 1997. For some insects, correspondence was quite close; for other insects, reliance on the Michigan sequence would have resulted in mistimed treatments. These data suggest that phenological indicators developed in one region may need to be evaluated on a case-by-case basis before they can be used with confidence in another region.
The tremendous diversity of ornamental plants, each with its own complement of insect pests, creates a logistical challenge for planning and implementing a successful pest management program. Insecticide applications must be timed precisely to maximize their effectiveness and minimize the number required. Improperly timed insecticide applications are expensive, environmentally detrimental, and result in customer dissatisfaction. Many insects are difficult to detect and monitor, further complicating the accurate timing of pesticide applications. Consequently, pesticide applications are frequently scheduled on a calendar-day basis. However, because of variation in patterns of degree-day accumulation from place to place and year to year, calendar-based scheduling is frequently inaccurate.
The use of ornamental plants as phenological indicators provides an alternative approach to predicting insect activity. Phenology is the study of recurring biological phenomena and their relationship to weather. Bird migration, hunting and gathering seasons, blooming of wildflowers and trees, and the seasonal appearance of insects are examples of phenological events that have been recorded for centuries (4,8). Because the development of both plants and insects is temperature dependent (6,12), plants may accurately track the environmental factors that affect insect development. Indeed, the use of plant phenology to predict insect activity is an old practice, with recorded observations that date back at least to the 18th century (7).
In recent years, two sets of phenological indicators have been published that demonstrate that phenological indicators can be useful tools for predicting the activity of insect pests. Donald A. Orton and Thomas L. Green published an extensive list of observations made in Illinois in their book Coincide (10). In addition, this author published a phenological sequence based on five years of data collected in central Michigan (5).
A critical but unresolved question is whether phenological indicators developed in one geographic region can be used accurately in another. The objectives of this report are:
During 1997, the phenology of 56 plant species and/or cultivars and 22 species of insects on or near the Ohio Agricultural Research and Development Center's Wooster campus were monitored (Table 1). For clarity, only common names are listed. To achieve standardization in nomenclature, common names of plants follow Dirr (3), and insect names are official common names as approved by the Entomological Society of America.
Plants were chosen to represent a range of blooming times from early March through late July. This time period corresponds with the activity of most of the important insect pests of ornamental plants. Four individuals of each species or cultivar were monitored. All individuals of a species were located either in uniform sun or shade, depending on the environment to which the species is adapted. Plants in microenvironments obviously altered by buildings, parking lots, and related areas were not used. Plants were monitored at least three times each week, and the dates of "first bloom" and "full bloom" were recorded. "First bloom" was defined as the date on which the first flower bud on the plant opened, revealing pistils and/or stamens, and "full bloom" as the date on which 95% of the flower buds have opened (i.e., one bud out of 20 has yet to open). These phenological events can be identified and recorded with precision.
The phenology of 22 insect pests with diverse life histories was also monitored in 1997, including defoliators, scales, gall formers, wood borers, and leafminers. As opposed to methods used to monitor plant phenology, which were designed to minimize variation in order to increase predictive power, sampling protocols for insects were designed to characterize the phenology of the entire population.
The Michigan phenological sequence developed at Dow Gardens in Midland consisted of 55 plant and 24 insect species, the phenology of which was monitored for five years (1985-1989) using the exact protocols described previously (5). Since the Michigan and Wooster phenological sequences have in common 40 plant and 14 insect species, they can be directly compared in order to test the accuracy of a sequence developed in one region (Michigan) for predicting insect activity in another (Wooster).
Table 1. Phenological sequence for Wooster, Ohio, in 1997 (insect names are indicated in bold type). | ||
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Species | Phenological Event | Date |
Silver Maple | first bloom | 7 March |
Corneliancherry Dogwood | first bloom | 15 March |
Red Maple | first bloom | 25 March |
Silver Maple | full bloom | 25 March |
Border Forsythia | first bloom | 30 March |
Manchu Cherry | first bloom | 2 April |
Korean Rhododendron | first bloom | 3 April |
Eastern Tent Caterpillar | egg hatch | 4 April |
Red Maple | full bloom | 4 April |
Star Magnolia | first bloom | 4 April |
Border Forsythia | full bloom | 4 April |
Corneliancherry Dogwood | full bloom | 4 April |
Sargent Cherry | first bloom | 5 April |
Norway Maple | first bloom | 6 April |
European Pine Sawfly | egg hatch | 15 April |
Inkberry Leafminer | adult emergence | 15 April |
Weeping Higan Cherry | first bloom | 20 April |
'Bradford' Callery Pear | first bloom | 21 April |
Rhododendron 'PJM' | first bloom | 21 April |
'Springsnow' Crabapple | first bloom | 22 April |
Allegheny Serviceberry | first bloom | 23 April |
Floweringquince | first bloom | 23 April |
Koreanspice Viburnum | first bloom | 26 April |
Allegheny Serviceberry | full bloom | 29 April |
Weeping Higan Cherry | full bloom | 29 April |
Rhododendron 'PJM' | full bloom | 1 May |
'Snowdrift' Crabapple | first bloom | 2 May |
'Donald Wyman' Crabapple | first bloom | 4 May |
Birch Leafminer | adult emergence | 5 May |
Japanese Flowering Crabapple | first bloom | 5 May |
Eastern Redbud | first bloom | 7 May |
Koreanspice Viburnum | full bloom | 7 May |
Gypsy Moth | egg hatch | 7 May |
Wayfaringtree Viburnum | first bloom | 10 May |
Sargent Crabapple | first bloom | 10 May |
'Coral Burst' Crabapple | first bloom | 11 May |
Tatarian Honeysuckle | first bloom | 12 May |
Ohio Buckeye | first bloom | 14 May |
'Springsnow' Crabapple | full bloom | 15 May |
'Snowdrift' Crabapple | full bloom | 15 May |
Common Lilac | first bloom | 15 May |
Honeylocust Plant Bug | egg hatch | 16 May |
'Donald Wyman' Crabapple | full bloom | 16 May |
'Coral Burst' Crabapple | full bloom | 18 May |
Imported Willow Leaf Beetle | adult emergence | 18 May |
Red Buckeye | first bloom | 18 May |
Sargent Crabapple | full bloom | 19 May |
Blackhaw Viburnum | first bloom | 19 May |
Red Chokeberry | first bloom | 19 May |
Eastern Redbud | full bloom | 19 May |
'Pink Princess' Weigela | first bloom | 20 May |
Wayfaringtree Viburnum | full bloom | 20 May |
Blackhaw Viburnum | full bloom | 22 May |
Japanese Flowering Crabapple | full bloom | 22 May |
Eastern Spruce Gall Adelgid | egg hatch | 22 May |
Pine Needle Scale | egg hatch | 22 May |
Vanhoutte Spirea | first bloom | 25 May |
Umbrella Magnolia | first bloom | 26 May |
Red Chokeberry | full bloom | 26 May |
Common Lilac | full bloom | 26 May |
Winter King Hawthorn | first bloom | 26 May |
Redosier Dogwood | first bloom | 27 May |
Tatarian Honeysuckle | full bloom | 28 May |
Holly Leafminer | adult emergence | 29 May |
Slender Deutzia | first bloom | 29 May |
Black Cherry | first bloom | 29 May |
Euonymus Scale | egg hatch | 30 May |
Lilac Borer | adult emergence | 30 May |
Pagoda Dogwood | first bloom | 30 May |
Ohio Buckeye | full bloom | 31 May |
Vanhoutte Spirea | full bloom | 1 June |
Scarlet Firethorn | first bloom | 1 June |
'Pink Princess' Weigela | full bloom | 2 June |
'Red Prince' Weigela | first bloom | 3 June |
Beautybush | first bloom | 3 June |
Black Cherry | full bloom | 3 June |
Lesser Peach Tree Borer | adult emergence | 4 June |
Winter King Hawthorn | full bloom | 5 June |
Redosier Dogwood | full bloom | 5 June |
Red Buckeye | full bloom | 7 June |
Sweet Mockorange | first bloom | 8 June |
Scarlet Firethorn | full bloom | 8 June |
Black Locust | first bloom | 9 June |
Umbrella Magnolia | full bloom | 9 June |
Common Ninebark | first bloom | 9 June |
Oystershell Scale | egg hatch | 9 June |
White Fringetree | first bloom | 9 June |
Pagoda Dogwood | full bloom | 9 June |
Bronze Birch Borer | adult emergence | 12 June |
Arrowwood Viburnum | first bloom | 12 June |
Mountain-laurel | first bloom | 12 June |
Black Locust | full bloom | 13 June |
White Fringetree | full bloom | 13 June |
American Holly | first bloom | 13 June |
Bumald Spirea | first bloom | 14 June |
Juniper Scale | egg hatch | 14 June |
Common Ninebark | full bloom | 15 June |
Potato Leafhopper | adult appearance | 16 June |
Arrowwood Viburnum | full bloom | 17 June |
American Holly | full bloom | 19 June |
'Red Prince' Weigela | full bloom | 20 June |
Washington Hawthorn | first bloom | 20 June |
Japanese Tree Lilac | first bloom | 21 June |
Sweetbay Magnolia | first bloom | 22 June |
American Elderberry | first bloom | 22 June |
Northern Catalpa | first bloom | 22 June |
Slender Deutzia | full bloom | 23 June |
Fall Webworm | first instars | 24 June |
Sweet Mockorange | full bloom | 24 June |
Sweetbay Magnolia | full bloom | 24 June |
Washington Hawthorn | full bloom | 24 June |
Mountain-laurel | full bloom | 25 June |
Northern Catalpa | full bloom | 25 June |
American Elderberry | full bloom | 28 June |
Littleleaf Linden | first bloom | 29 June |
Spruce Bud Scale | egg hatch | 30 June |
Bumald Spirea | full bloom | 2 July |
Japanese Beetle | adult emergence | 2 July |
Panicled Goldenraintree | first bloom | 7 July |
Rosebay Rhododendron | first bloom | 7 July |
Littleleaf Linden | full bloom | 7 July |
Peach Tree Borer | adult emergence | 8 July |
Rosebay Rhododendron | full bloom | 20 July |
Panicled Goldenraintree | full bloom | 28 July |
Magnolia Scale | first instars | 3 August |
The phenological sequence observed in Wooster, Ohio, in 1997 is presented in Table 1. The sequence of occurrence is much more important than the dates, which are only included to provide perspective, and which will vary dramatically with year-to-year variation in degree-day accumulation. During the cool spring of 1997, plant and insect phenology was substantially delayed relative to "normal" years.
It is important to emphasize that any conclusions drawn from this sequence are preliminary, as they are based on only one year of data. Because consistency in the sequence is essential if the sequence is to be useful for predicting insect activity, this study will continue for several more years to test this critical assumption. However, in the five-year Michigan study, there was substantial year-to-year variation in degree-day accumulation, yet little variation in the order in which plants bloomed and insects appeared from one year to the next.
When a phenological sequence of plants can be shown to correspond with the appearance of insects, pest managers can use the easily monitored plant sequence as a "biological clock" to anticipate the order and the time when pests reach vulnerable stages. This can greatly facilitate the logistics of serving many clients with a variety of problems. For example, one can see from Table 1 that when common lilac was in full bloom:
Again, it must be emphasized that this first year of data should be considered preliminary. It remains to be seen if this sequence proves valid as the study continues.
A useful attribute of phenological sequences is that they can be readily expanded and customized. Once the basic sequence is in place, any new plants or pests can be added as the need occurs. For example, if a pest manager has made a particularly successful treatment, any plants in bloom at the time can be noted, added to the sequence, and used to duplicate the timing in future years.
This critical question must be answered before published phenological sequences can be used with confidence outside the regions in which they were developed. To the degree that phenological relationships do not hold across regions, it will be necessary to develop localized sequences. Table 2 can be used to evaluate the accuracy of the Michigan plant sequence for predicting insect phenology in Ohio. The order in which phenological events occurred in Ohio in 1997 is compared with the average order in which they occurred in Michigan from 1985-1989. The magnitude of any disparity in the order in which a particular event occurred in the two sequences is also shown. A positive value indicates that the event occurred earlier in the sequence in Michigan that it did in Ohio, while a negative value indicates the opposite.
The correspondence in the flowering sequences of the plants common to both studies was quite close. Flowering of silver maple occurred first in both locations, followed by flowering of corneliancherry dogwood. Red maple flowered just before border forsythia in Ohio, and just after it in Michigan; first bloom of Norway maple was the ninth event in both sequences. In fact, in no case did the order of flowering by any species vary by more than the five places between the two sequences. As more plant species are included in the sequence, the chance of being led astray by one species that departs from the pattern will diminish.
The sequence of insect phenological events did not correspond as closely. Egg hatch of eastern tent caterpillar occurred before first bloom of red maple, border forsythia, and Korean rhododendron in Michigan, but after their first bloom in Ohio. Some of the discrepancies were substantial enough that any pesticide applications on which they had been based would have been mistimed and probably inefficacious. For example, egg hatch of oystershell scale closely followed first bloom of Vanhoutte spirea in Michigan, but did not occur until more than two weeks after first bloom in Ohio. There was even less correspondence in the phenology of euonymus scale egg hatch, which occurred just after first bloom of black cherry in Ohio, but not until after first bloom of common ninebark in Michigan. The much closer correspondence in other species including birch leafminer, pine needle scale, bronze birch borer, and juniper scale suggests that phenological sequences may have regional generality, at least in some cases.
A number of factors may affect the accuracy of plant phenological indicators as predictors of insect activity. The assumption that a given phenological correlation will occur in different climatic regions requires that all organisms included in the correlation have the same upper and lower temperature thresholds for development, as well as the same developmental responses to changing temperature. These traits are known to vary widely among both plants and insects (6,12).
Origin, or provenance, of indicator plants can affect their accuracy as phenological indicators. Both elevational and latitudinal gradients can affect plant phenology, as different provenances are adapted to different day-length and temperature regimes (12). Furthermore, the same genotype may display different phenological patterns when planted in different geographic regions (9). Substantial geographic variation has also been documented in the phenological patterns of insects (1, 2, 13).
Data from the Michigan study indicate that insects that overwinter in the soil, such as the imported willow leaf beetle and white pine weevil, are less accurately predicted from year-to-year than insects that overwinter on aerial plant parts exposed to the same ambient conditions as flower buds. The phenology of insects overwintering in the soil may be influenced substantially by a number of factors not having as strong an effect on the phenology of flower buds, including vegetation, snow cover, and soil moisture (11).
Phenological indicators will be more accurate in the region in which they are developed because these factors are less variable within a region than among different regions. Furthermore, at any given location the weather is more similar from one year to the next than is the weather from one location to another. The results of this study suggest that while some phenological indicators may be regionally robust, others are not. Hence, phenological indicators developed in one region will need to be evaluated on a case-by-case basis before they can be used with confidence in another region. None the less, the use of plant phenological indicators for timing pest activity holds tremendous potential for improving the effectiveness of integrated pest management programs in the landscape.
Table 2. Comparison of a phenological sequence for Wooster, Ohio, in 1997 with a phenological sequence from Midland, Michigan, from 1985‚1989 (insect names are indicated by bold type). | ||||
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Species | Event | Order in Ohio | Order in Michigan | Disparity in Rank |
Silver Maple | first bloom | 1 | 1 | 0 |
Corneliancherry Dogwood | first bloom | 2 | 2 | 0 |
Red Maple | first bloom | 3 | 4 | -1 |
Border Forsythia | first bloom | 4 | 5 | -1 |
Manchu Cherry | first bloom | 5 | 8 | -3 |
Korean Rhododendron | first bloom | 6 | 7 | -1 |
Eastern Tent Caterpillar | egg hatch | 7 | 3 | 4 |
Star Magnolia | first bloom | 8 | 6 | 2 |
Norway Maple | first bloom | 9 | 9 | 0 |
Weeping Higan Cherry | first bloom | 10 | 14 | -4 |
'Bradford' Callery Pear | first bloom | 11 | 12 | -1 |
'PJM' Rhododendron | first bloom | 12 | 11 | 1 |
Allegheny Serviceberry | first bloom | 13 | 15 | -2 |
Floweringquince | first bloom | 14 | 16 | -2 |
Koreanspice Viburnum | first bloom | 15 | 19 | -4 |
'Snowdrift' Crabapple | first bloom | 16 | 21 | -5 |
Birch Leafminer | adult emergence | 17 | 18 | -1 |
Japanese Flowering Crabapple | first bloom | 18 | 20 | -2 |
Eastern Redbud | first bloom | 19 | 17 | 2 |
Gypsy Moth | egg hatch | 20 | 13 | 7 |
Wayfaringtree Viburnum | first bloom | 21 | 24 | -3 |
'Coral Burst' Crabapple | first bloom | 22 | 25 | -3 |
Tatarian Honeysuckle | first bloom | 23 | 26 | -3 |
Ohio Buckeye | first bloom | 24 | 23 | 1 |
Common Lilac | first bloom | 25 | 22 | 3 |
Willow Leaf Beetle | adult emergence | 26 | 10 | 16 |
Blackhaw Viburnum | first bloom | 27 | 29 | -2 |
Doublefile Viburnum | first bloom | 28 | 30 | -2 |
Cooley Spruce Gall Adelgid | egg hatch | 29 | 27 | 2 |
Pine Needle Scale | egg hatch | 30 | 28 | 2 |
Vanhoutte Spirea | first bloom | 31 | 33 | -2 |
'Winter King' Hawthorn | first bloom | 32 | 37 | -5 |
Black Cherry | first bloom | 33 | 32 | 1 |
Euonymus Scale | egg hatch | 34 | 43 | -9 |
Lilac Borer | adult emergence | 35 | 31 | 4 |
Pagoda Dogwood | first bloom | 36 | 35 | 1 |
Beautybush | first bloom | 37 | 38 | -1 |
Lesser Peachtree Borer | adult emergence | 38 | 36 | 2 |
Black Locust | first bloom | 39 | 39 | 0 |
Common Ninebark | first bloom | 40 | 41 | -1 |
Oystershell Scale | egg hatch | 41 | 34 | 7 |
White Fringetree | first bloom | 42 | 40 | 2 |
Bronze Birch Borer | adult emergence | 43 | 42 | 1 |
Mountain-laurel | first bloom | 44 | 44 | 0 |
Juniper Scale | egg hatch | 45 | 45 | 0 |
Washington Hawthorn | first bloom | 46 | 46 | 0 |
Japanese Tree Lilac | first bloom | 47 | 47 | 0 |
American Elderberry | first bloom | 48 | 50 | -2 |
Northern Catalpa | first bloom | 49 | 48 | 1 |
Littleleaf Linden | first bloom | 50 | 52 | -2 |
Spruce Bud Scale | egg hatch | 51 | 54 | -3 |
Panicled Goldenraintree | first bloom | 52 | 53 | -1 |
Rosebay Rhododendron | first bloom | 53 | 49 | 4 |
Peachtree Borer | adult emergence | 54 | 51 | 3 |
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10. Orton, D. A. and T. L. Green. 1989. Coincide. Plantsmen's Publications, Flossmoor, Il.
11. Raffa, K. F. and D. W. A. Hunt. 1989. Microsite and interspecific interactions affecting emergence of root-infesting pine weevils (Coleoptera: Cur-culionidae) in Wisconsin. Annals of the Entomological Society of America 82:438-445.
12. Rathcke, B. and E. L. Lacey. 1985. Phenological patterns of terrestrial plants. Annual Review of Ecology and Systematics 16:179-214.
13. Tauber, C. A. and M. J. Tauber. 1981. Insect seasonal cycles: genetics and evolution. Annual Review of Ecology and Systematics 12:281-308.