Animal Diversity Web

Diversity

Members of the family Geometridae are commonly called geometer moths, and their larvae are commonly called loopers or inchworms, due to their unique form of locomotion. Geometer moth larvae typically have only two pairs of prolegs at their posterior ends, in contrast to the five pairs that are common in most moth families.

Geometer moths are abundant around the world and are widely known as one of the three most speciose families in the order Lepidoptera. The most recent estimates suggest there are nearly 24,000 species in nine subfamilies worldwide. Many of the discovered species are endemic to North or South America.

Adult geometer moths are classically characterized by wing position and shape when at rest. They tend to hold their wide forewings flat against the ground, away from their bodies. Furthermore, their wings are usually patterned with mottled splotches or wavy lines. geometer moths are generally intermediate in size, with wingspans between 15 and 50 mm. Despite these similarities, species in the family Geometridae are otherwise morphologically diverse. Some geometer moths resemble species in the moth family Erebidae. (Bodner, et al., 2010; McLeod, 2022; Murillo-Ramos, et al., 2019)

Geographic Range

Geometer moths occur in all biogeographic regions except for Antarctica. Because the family Geometridae is so speciose, there are both native and introduced species across the globe. (Bodner, et al., 2010; Fauske, 2002; Lotts, et al., 2023; McLeod, 2022; Murillo-Ramos, et al., 2019)

• Biogeographic Regions
• nearctic
  • native
• palearctic
  • native
• oriental
  • native
• ethiopian
  • native
• neotropical
  • native
• australian
  • native
• oceanic islands
  • native

• Other Geographic Terms
• cosmopolitan

Habitat

The family Geometridae consists of around 24,000 known species spread across the globe. Different species occupy different habitats depending on their geographic distribution. Geometer moths occupy tropical, subtropical, temperate, and polar zones at a wide range of elevations, including montane and coastal environments. The larvae of many species primarily eat leaves of woody plants and thus many species are associated with forests and scrublands. Despite their preference for woody plants, most geometer moths have flexible diets and are also capable of persisting in cities and agricultural areas. (Axmacher, et al., 2009; Beck, et al., 2002; Özdemir, 2019)

• Habitat Regions
• temperate
• tropical
• polar

• Terrestrial Biomes
• tundra
• taiga
• desert or dune
• savanna or grassland
• chaparral
• forest
• rainforest
• scrub forest
• mountains

• Wetlands
• marsh
• swamp
• bog

• Other Habitat Features
• urban
• suburban
• agricultural

Systematic and Taxonomic History

Geometer moths are considered the second largest family within the order Lepidoptera. There are nine subfamilies that collectively comprise over 2,000 genera, with nearly 24,000 currently recognized species worldwide. However, it is likely that these numbers underestimate the true diversity in the family Geometridae and some species have not yet been described or distinguished from morphologically similar groups. Further genetic research is required to identify cryptic species and better describe the true diversity of geometer moths. (Murillo-Ramos, et al., 2019; Rajaei, et al., 2022)

• Synapomorphies

  • Trifid venation of cubitus
  • Tympana present

Physical Description

Adult geometer moths are typically small or intermediate in body size. Adults are characterized by slender bodies and broad wings, which many species hold outspread and flat against surfaces when at rest. Many geometer species have disruptive coloration. Their wings and bodies are typically brown, grey, or green with mottling or continuous wavy lines along their forewings and hindwings. Geometer moths have paired tympana, located ventrally on their first abdominal segment.

Geometer moths exhibit sexual dimorphism in the morphology of their antennae. Generally, females have thin antennae (filiform) and males have feathery antennae (pectinate). In some species, females have extremely reduced wings and are incapable of flight.

Geometer moth larvae, commonly called inchworms or loopers, are distinguishable from most other larval lepidopterans (order Lepidoptera). Most lepidopteran larvae have five pairs of forelegs from their posterior ends to their midbodies, whereas inchworms have only two or three pairs of prolegs near their posterior ends. Because of this morphological difference, inchworms exhibit a unique form of locomotion distinct from the undulating movement of most lepidopteran larvae. Inchworms move by anchoring with their thoracic legs and bringing their prolegs forward, just behind their thoracic legs, forming an arch in the middle of their bodies. They then anchor with their prolegs and extend the anterior part of their body forward before anchoring again with their thoracic legs. Inchworms are typically slender and often have coloration and patterns that resemble twigs or the stems of leaves. (Lotts, et al., 2023; McLeod, 2022; Meyer and Orr, 2015)

• Other Physical Features
• ectothermic
• bilateral symmetry

• Sexual Dimorphism
• sexes alike

Development

Geometer moths undergo complete metamorphosis; larvae hatch from eggs and go through a pupal stage before reaching adulthood. Geometer moths are often bivoltine, with two distinct generations hatching in early spring and late summer, respectively. Females lay their eggs on leaves, branches, or bark crevices in late spring or in late fall. Eggs that are laid in late summer or fall overwinter as eggs and hatch in spring. In contrast, eggs that are laid in spring or early summer either hatch, mature, and reproduce before winter or pupate and enter diapause during winter, only emerging as adults during spring.

Adult female geometer moths lay their eggs, either singly or in clusters, on leaf surfaces, branches, or in bark crevices. Larval geometer moths typically feed on the leaves of woody plants, though diets vary between Geometridae species. They then descend to the ground and pupate in the soil or underneath leaf litter. Geometer moths exhibit significant phenotypic plasticity depending on environmental conditions, such as ambient temperature, day length, and population density. Generally, growth rates are faster and body size is smaller in individuals that undergo direct development, whereas individuals that enter diapause develop slower and have larger body sizes as larvae and pupae. Higher ambient temperatures and longer day length tend to promote faster growth rates and direct development.

Diet composition also has an impact on development, with higher quality foods (e.g., young leaves) increasing body size and fecundity. In many species, the timing of larval emergence in spring coincides with the emergence of young leaves on preferred host plant species. Furthermore, diet can impact the expression of morphological traits. For example, one species of emerald moth, Nemoria arizonaria, mimics different features of its host plant depending on its diet: individuals that eat oak catkins develop features that mimic the catkin structure, whereas those that eat oak leaves mimic small twigs. (Choi and Kim, 2014; Drooza, 1970; Greene, 1996; Haukioja, et al., 1988; Kivelä, et al., 2012; Lotts, et al., 2023; Välimäki, et al., 2013; Wint, 1983)

• Development - Life Cycle
• metamorphosis
• diapause

Reproduction

Geometer moths are typically polygynous, but occasionally exhibit polygynandrous behavior. For example, a 1974 study on hemlock loopers (Lambdina fiscellaria) found that females typically receive a spermatophore from only one male, but have been recorded to receive spermatophores from up to three males.

Adult geometer moths are typically short-lived (approximately 5 - 20 days) and mate within a few days after pupal eclosion. Recently eclosed females select a location, often among vegetation, and release pheromones to attract males. In many geometer moth species males have large, pennate antennae that help them detect and follow pheromone trails to reproductive females. Such antennae are particularly pronounced in species with non-volant females. During copulation, males deposit a bundle of sperm, called a spermatophore, which females store and use to fertilize their eggs. (Choi and Kim, 2014; Javoiš, et al., 2019; Ostaff, et al., 1974; Yang, et al., 2011)

• Mating System
• polygynous
• polygynandrous (promiscuous)

Within the family Geometridae females vary widely in adult body size and initial fecundity. Females of some species, such as winter moths (Operophtera brumata) and fall cankerworms (Alsophila pometaria), have enlarged abdomens and wings that are significantly reduced or entirely absent. In these species, females emerge from their pupal stage with a large number of eggs and do not feed as adults. Such species are capital breeders, using stored energy from their larval period to gestate and lay eggs. Alternatively, some geometer moth species (called income breeders) are volant and forage as adults, with foraging success being positively related to fecundity.

In capital breeding geometer moth species, adults are typically active in cooler weather during spring or fall, whereas income breeders are mostly active in summer. Furthermore, species that are capital breeders tend to show indiscriminate oviposition, laying their eggs on non-host plants and on plant surfaces other than leaves. In contrast, most income breeders are more selective, laying their eggs on the leaves of preferred host plants. Female geometer moths lay the majority of their eggs, either singly or in clusters, shortly after copulation. The number of eggs that geometer moths lay varies depending on the species and individual body mass, as well as environmental factors such as temperature, food quality and availability, and population density. (Choi and Kim, 2014; Davis, et al., 2016; Javoiš, et al., 2019; Steinbauer, 2005; Tammaru, et al., 1995; Werner, 2012)

• Key Reproductive Features
• semelparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• fertilization
  • internal
• oviparous
• sperm-storing

Geometer moths exhibit very little parental investment. In many Geometridae species, females lay their eggs on leaf surfaces of host plants so that newly hatched larvae have access to food sources. However, larvae are completely independent upon hatching. Males exhibit no parental investment beyond the act of mating. Adults typically die within 20 days after emerging from their pupal stage. (Choi and Kim, 2014; Davis, et al., 2016)

• Parental Investment
• no parental involvement
• pre-hatching/birth
  • provisioning
    • female

Lifespan/Longevity

Geometer moths typically live less than a year, although individual longevity varies depending on the time of year that they emerge from eggs. Larvae that hatch in spring or early summer often complete metamorphosis and die by early fall. In other cases, individuals overwinter as eggs, larvae, or pupae and do not complete metamorphosis until the following spring. Adult geometer moth lifespans are typically around 5 to 9 days, but individuals have been recorded to live for more than 30 days. Across the Geometridae family, larger-bodied species tend to live slightly longer. (Holm, et al., 2016; Holm, et al., 2022; Kivelä, et al., 2012)

Behavior

Geometer moth adults are short-lived, so they mainly exhibit reproductive behaviors after eclosing from pupae. Adult females rest on vegetation shortly after eclosion and emit pheromones to attract males. Males have feather-like (pectinate) antennae, which are highly sensitive to female pheromones and help males locate receptive females. Following successful copulation, females seek oviposition sites whereas males continue to seek other mates. Females generally select oviposition sites that provide suitable foraging opportunities for larvae. Geometer moths are mostly nocturnal and exhibit erratic flight patterns when searching for mates or oviposition sites. Females of some species have reduced wings or lack wings entirely, so they are less mobile and tend to be less selective about where they lay their eggs.

Geometer moth larvae have a distinctive form of locomotion. They begin by bringing their rear legs up to their forelegs, forming an arch in the middle of their bodies, and then straighten their bodies to move forward. In temperate climates, geometer moths may enter diapause and overwinter as larvae or pupae before completing metamorphosis. Although geometer moths are solitary, larvae can reach high population densities in areas with high levels of host plant species. Larvae spend the majority of their time foraging on host plants. Several species endemic to the Hawaiian islands are carnivorous ambush predators. They camouflage as small twigs on tree branches and strike at small insects that happen to approach. (Lotts, et al., 2023; McLeod, 2022; Montgomery, 1983; Tammaru, et al., 1995)

• Key Behaviors
• arboreal
• flies
• nocturnal
• motile
• hibernation
• solitary

Communication and Perception

Geometer moths perceive their environment using chemical, visual, tactile, and auditory stimuli. The majority of geometer moth species are exclusively nocturnal as adults. They have well-developed night vision, which they use to navigate while flying. Geometer moths also have tympana located ventrally on each side of their first abdominal segments, which are sensitive to frequencies that bats commonly produce when echolocating. These tympana help geometer moths avoid predation by sympatric bat species. Adult geometer moths also perceive chemical cues in their environment. For example, taste receptors on their legs help them detect chemicals found in their host plants, which is beneficial to geometer moths when foraging or selecting oviposition sites. Larvae are also capable of discerning high-quality host plants based on chemical signals. Furthermore, at least some species adjust the expression of chemicals in their body depending on the plants they eat.

Geometer moths also use chemical cues to communicate with each other. Research suggests that larvae may regurgitate chemicals that restrict the growth of conspecifics that come in direct contact. For adults, chemical communication is critical to reproductive success. Adult males have pectinate antennae that help them follow pheromone trails produced by reproductively active females. During copulation, geometer moths also rely on tactile stimuli to align their genitalia for the successful transfer of spermatophores. Caterpillars may also respond to physical stimuli, (Akino, 2005; Gruys, 1970; McLeod, 2022; Meyer-Rochow and Lau, 2008; Pöykkö, 2006; Rydell, et al., 1997; Wint, 1983)

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Other Communication Modes
• pheromones

• Perception Channels
• visual
• tactile
• ultrasound
• chemical
• electric

Food Habits

The vast majority of geometer moth species are herbivorous. Some species are dependent on one host plant species (monophagous) or a select few host species (oligophagous), while other species are more flexible and use multiple types of plants as hosts (polyphagous). Members of the family Geometridae are distributed widely throughout the globe, with species native to every continent except Antarctica. As a result, geometer moth species feed on a wide variety of plant species, including gymnosperms, angiosperms, and pteridophytes. Most geometer moth species use woody shrubs or trees as host plants, although some species feed on herbaceous plants or, in some cases, lichens.

Geometer moth larvae typically eat leaves, though they may also eat other plant structures, such as stems, petals, pollen, seeds, or leaf buds. The larvae of several native Hawaiian species in the genus Eupithecia are carnivorous. These species camouflage as small twigs on branches, with the anterior portion of their bodies held aloft, and opportunistically strike at insects that fly or walk nearby. Adult geometer moths eat less than larvae, instead spending most of their time searching for mates. Some species do not eat as adults, instead using stored energy from their larval phase to emerge from pupae with a large number of developed eggs. The geometer moth species that do eat as adults typically consume liquids, such as pollen or tree sap. In these species, the amount of food acquired during adulthood is positively correlated with the number of eggs laid. (Bodner, et al., 2010; Montgomery, 1983; Robinson, et al., 2010)

• Primary Diet
• carnivore
  • insectivore
• herbivore
  • folivore
  • granivore
  • nectarivore

Predation

Geometer moths serve as prey for various other arthropods, such as true bugs (order Hemiptera) and spiders (order Araneae), as well as vertebrate predators, such as small mammals, birds, reptiles, and amphibians. Because geometer moths are globally distributed, specific predators vary depending on the geographic range of any given geometer moth species.

Many geometer moths have cryptic coloration, both as larvae and adults. Caterpillars are often green or brown and tend to resemble small twigs or the stems of leaves. Adults are generally brown, grey, or green, with patterns along their wings that disrupt their silhouettes. Geometrid caterpillars are capable of detecting and distinguishing substrate vibrations created by potential predators. They exhibit different anti-predator behaviors depending on the threat. In some cases, as with large predators like birds, caterpillars remain still until the threat is gone. In other cases, as with predatory insects, they use silk to suspend themselves away from vegetation or drop to the ground. There are also instances where caterpillars will show aggression towards potential predators, attacking them with their mouthparts. (Berndt, et al., 2012; Bodner, et al., 2010; Castellanos, 2010; Castellanos and Barbosa, 2006; Connahs, et al., 2009; Soares, et al., 2009; Tanhuanpää, et al., 2001a; Tanhuanpää, et al., 2001b)

• Anti-predator Adaptations
• cryptic

Ecosystem Roles

Geometer moth caterpillars are herbivores and can be detrimental to the health of their host plants. This is especially true when populations reach high densities. In such outbreak conditions, caterpillars can defoliate large stretches of vegetation, and there is evidence that this activity can influence local succession patterns. Some geometer moth species are monophagous, meaning the extent of these effects are limited to one host plant species. However, polyphagous geometer moth species have the potential to impact multiple plant species.

Some geometer moth species consume flower nectar as adults, making them important pollinators for plant species within their range. However, the extent to which geometer moths benefit plants through pollination depends on the level of herbivory that larvae exhibit on those same plants.

Geometer moths are susceptible to parasitoids such as tachinid flies (family Tachinidae) and wasps from the families Braconidae, Ichneumonidae, Chalcididae, and Eulophidae. In a study of Geometer life histories in South America, only 32% of caterpillars collected were successfully raised to adulthood and 25% of premature mortality was attributed to parasitoidism. Most geometer moths are likely susceptible to parasitoidism , although research suggests that the number of parasitoids and the frequency of parasitoid attacks may be influenced by habitat type. (Guthrie, et al., 2012; Hahn and Brühl, 2016; Karlsen, et al., 2013; MacGregor, et al., 2015; Tanhuanpää, et al., 2001a; Westerbergh, 2004)

• Ecosystem Impact
• pollinates

Economic Importance for Humans: Positive

There is limited information regarding the positive economic impacts of geometer moths. Some species act as important plant pollinators, although the role they play in pollinating crop plants is not well studied. Geometer moth larvae and adults may also serve as an important source of food for other fauna. However, the economic benefits of geometer moth activity have not been quantified. (Hahn and Brühl, 2016; MacGregor, et al., 2015)

Economic Importance for Humans: Negative

Geometer moths can have negative economic impacts on crop productivity, as larval activity can result in significant plant defoliation. Certain species of geometer moths feed on common crop plants, including tea (Camellia sinensis), wild blueberry (Vaccinium angustifolium), and eucalyptus plants (genus Eucalyptus). Furthermore, some geometer moth species use deciduous or coniferous trees as host plants, and thus can have large effects on forestry and silviculture.

The large-scale defoliation associated with geometer moth outbreaks can substantially increase host plant mortality. High rates of woody plant mortality can impact forest structure and community composition. Consequently, geometer moths can have indirect, cascading effects on ecosystem functions and the local distribution of land cover types. Furthermore, worsening global climate change may result in geometer moth range shifts, in which case new areas would be impacted by defoliation. (Caldwell, et al., 2005; Ding, et al., 2022; Oliveira, et al., 2008; Ramanaidu, et al., 2011; Roy, et al., 2017; Tuffen, et al., 2019; Wint, 1983)

• Negative Impacts
• crop pest

Conservation Status

Geometer moths exhibit a range of life history traits, depending on the species. Polyphagous species with wide geographic distributions tend to be more resilient to environmental changes and habitat loss. In some cases, such species may benefit from climate change, dispersing to new areas as conditions become suitable. Alternatively, geometer moth species that are oligophagous or have restricted ranges are at higher risk of population decline in the future. Perhaps one of the largest threat to such geometer moths is habitat loss due to anthropogenic activity. However, these effects are likely exacerbated by climate change and the introduction of invasive species, such as parasitoid flies and competitors for similar resources. (Ding, et al., 2022; McQuillan, 2004; Terblanche, 2018; Wagner, 2012; Waring, 2004)

• IUCN Red List [Link]

  Not Evaluated

Contributors

Galen Burrell (author), Special Projects.

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