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Abstract
The focus of this study was to reconstruct a phylogenetic hypothesis for the moth subfamily Arctiinae (tiger moths, woolly bears) to investigate the evolution of larval and adult pharmacophagy of pyrrolizidine alkaloids (PAs) and the pathway to PA chemical specialization in Arctiinae. Pharmacophagy, collection of chemicals for non-nutritive purposes, is well documented in many species, including the model species Utetheisa ornatrix L. A total of 86 exemplar ingroup species representing tiger moth tribes and subtribes (68 genera) and nine outgroup species were selected. Ingroup species included the most species-rich generic groups to represent the diversity of host-plant associations and pharmacophagous behaviors found throughout Arctiinae. Up to nine genetic markers were sequenced: one mitochondrial (COI barcode region), one nuclear rRNA (D2 region, 28S rRNA), and seven nuclear protein-coding gene fragments: elongation factor 1-α protein, wingless, ribosomal protein subunit S5, carbamoylphosphate synthase domain regions, glyceraldehyde-3-phosphate dehydrogenase, isocitrate dehydrogenase and cytosolic malate dehydrogenase. A total of 6984 bp was obtained for most species. These data were analyzed using model-based phylogenetic methods: maximum likelihood (ML) and Bayesian inference (BI). Ancestral pharmacophagous behaviors and obligate PA associations were reconstructed using the resulting Bayes topology and Reconstructing Ancestral States in Phylogenies (RASP) software. Our results corroborate earlier studies on the evolution of adult pharmacophagous behaviors, suggesting that this behavior arose multiple times and is concentrated in the phaegopterine-euchromiine-ctenuchine clade (PEC). Our results suggest that PA specialization may have arisen early in the phylogeny of the subfamily and that facultative larval pharmacophagous behaviors are the derived condition.
Highlights
Arctiinae are a charismatic moth lineage with a complex evolutionary relationship with plant and fungal chemistries
Both aposematic adults (Figure 1A) and larvae (Figure 1B–D) typically harbor endogenous biogenic amines, like histamines, which are often supplemented with secondary compounds acquired from larval hosts or through pharmacophagy - feeding on plants to obtain chemicals rather than nutrients [2]
Field locations where permits were required with relevant contact information are as follows: USA: Big Cypress National Preserve, FL: collecting and destructive sampling permits were obtained through National Parks Services (Jennifer Stafford and Steve Schulze); COSTA RICA: permits were obtained through the Organization for Tropical Studies (Francisco Campos Rivera); RUSSIAN FEDERATION: Primorsky Krai, permits obtained through the Far Eastern Branch of the Russian Academy of Science
Summary
Arctiinae (tiger moths and woolly bears) are a charismatic moth lineage with a complex evolutionary relationship with plant and fungal chemistries. Comprised of approximately 11,000 species [1], this cosmopolitan group is well known among ecologists and evolutionary biologists for the evolution of bright coloration and spectacular adult mimicry of wasps, beetles and unpalatable moths and butterflies (Figure 1). Both aposematic adults (Figure 1A) and larvae (Figure 1B–D) typically harbor endogenous biogenic amines, like histamines, which are often supplemented with secondary compounds acquired from larval hosts or through pharmacophagy - feeding on plants to obtain chemicals rather than nutrients [2]. Adult pharmocophagy has been documented in other Lepidoptera, notably Danainae (Nymphalidae) [2]; larval pharmacophagy has only been documented in Arctiinae
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