Carla M. Penz

Vertical distribution, flight behaviour and evolution of wing morphology in Morpho butterflies.

1. Flight is a key innovation in the evolution of insects that is crucial to their dispersal, migration, territoriality, courtship and predator avoidance. Male butterflies have characteristic territoriality and courtship flight behaviours, and females use a characteristic flight behaviour when searching for host plants. This implies that selection acts on wing morphology to maximize flight performance for conducting important behaviours among sexes. 2. Butterflies in the genus Morpho are obvious components of neotropical forests, and many observations indicate that they show two broad categories of flight behaviour and flight height. Although species can be categorized as using gliding or flapping flight, and flying at either canopy or understorey height, the association of flight behaviour and flight height with wing shape evolution has never been explored. 3. Two clades within Morpho differ in flight behaviour and height. Males and females of one clade inhabit the forest understorey and use flapping flight, whereas in the other clade, males use gliding flight at canopy level and females use flapping flight in both canopy and understorey. 4. We used independent contrasts to answer whether wing shape is associated with flight behaviour and height. Given a single switch to canopy habitation and gliding flight, we compared contrasts for the node at which the switch to canopy flight occurred with the distribution of values in the two focal clades. We found significant changes in wing shape at the transition to canopy flight only in males, and no change in size for either sex. A second node within the canopy clade suggests that other factors may also be involved in wing shape evolution. Our results reinforce the hypothesis that natural selection acts differently on male and female butterfly wing shape and indicate that the transition to canopy flight cannot explain all wing shape diversity in Morpho. 5. This study provides a starting point for characterizing evolution of wing morphology in forest butterflies in the contexts of habitat selection and flight behaviour. Further, these observations suggest that exploring wing shape evolution for canopy and understorey species in other insects may help understand the effects of habitat destruction on biological diversity.

Phylogenetic Analysis of Morpho Butterflies (Nymphalidae, Morphinae): Implications for Classification and Natural History

Abstract The classification of butterflies in the widely recognized genus Morpho previously used subgenera that were assumed to constitute natural species groups. Cladistic analysis of 120 characters provided a well-resolved tree showing that some subgenera do not constitute monophyletic groups. This study supported some traditional taxonomic species groupings, but rejected the concept of subgenera for Morpho. Therefore, we formally redefined the genus to be consonant with the assumptions of phylogenetic classification. Predictions about Morpho life histories, the correlation of color pattern and flight behavior with vertical flight height, and the evolution of sexual dimorphism are discussed in light of our phylogeny.

Adult and early-stage characters of Brassolini contain conflicting phylogenetic signal (Lepidoptera, Nymphalidae)

This study examines the contribution of early‐stages and adult characters to the reconstruction of the phylogeny of Brassolini butterflies. Parsimony analyses used both equal weights and implied weights, and a series of analyses were performed. First, we analysed adult and early‐stages partitions independently and in combination for a subset of 27 species; in these cases the matrices were mostly complete. Whereas the adult partition alone produced a topology that was well resolved and congruent with previous studies, the early‐stages partition produced a poorly resolved tree under equal weights. Furthermore, implied weights produced a well‐resolved early‐stages topology that differed significantly from the adult topology. When both partitions were combined for 27 species, implied weights yielded a topology that resembled the adult tree except for the positions of Bia and Penetes, but statistical node support was generally lower. This suggests that stochastic noise increased when early‐stage characters were added to the adult partition, but the combined partitions topology was not statistically different from that based on adult characters alone. Second, given that preserved early stages are not as readily available as adults, we analysed a matrix including 45 species in which early‐stage data were missing for 18 species, and compared the topology to that produced by the adult partition alone. Results were similar to the analyses including fewer species; the combined partitions tree was similar to that from the adult partition except for the position of Bia and Penetes. We compare our findings to other genus‐level phylogenetic studies within Lepidoptera that have also used early‐stages and adult characters.

Evaluating the monophyly and phylogenetic relationships of Brassolini genera (Lepidoptera, Nymphalidae)

Abstract This study uses 80 morphological characters and cladistic analysis to evaluate the monophyly and phylogenetic relationships of 18 genera that constitute the butterfly tribe Brassolini. Most characters derive from genitalia, confirming previous generic definitions based mainly on wing characters, and showing that 16 of 18 genera are monophyletic. Mimoblepia Casagrande, syn.n. was subsumed within Opoptera Aurivillius to address the paraphyly of the latter, but resolution of the status of Aponarope Casagrande requires further study. The results suggest that the Brassolini includes six suprageneric groups/clades. Although this study verifies some genus‐level relationships put forward over 100 years ago, some new hypotheses of relationships are proposed. Tracing larval host plant use onto the Brassolini phylogeny indicates that species in this tribe retain the use of Arecaceae and Poaceae from their ‘satyroid’ ancestors.

Cruising the rain forest floor: butterfly wing shape evolution and gliding in ground effect

Flight is a key innovation in the evolutionary success of insects and essential to dispersal, territoriality, courtship and oviposition. Wing shape influences flight performance and selection likely acts to maximize performance for conducting essential behaviours that in turn results in the evolution of wing shape. As wing shape also contributes to fitness, optimal shapes for particular flight behaviours can be assessed with aerodynamic predictions and placed in an ecomorphological context. Butterflies in the tribe Haeterini (Nymphalidae) are conspicuous members of understorey faunas in lowland Neotropical forests. Field observations indicate that the five genera in this clade differ in flight height and behaviour: four use gliding flight at the forest floor level, and one utilizes flapping flight above the forest floor. Nonetheless, the association of ground level gliding flight behaviour and wing shape has never been investigated in this or any other butterfly group. We used landmark-based geometric morphometrics to test whether wing shapes in Haeterini and their close relatives reflected observed flight behaviours. Four genera of Haeterini and some distantly related Satyrinae showed significant correspondence between wing shape and theoretical expectations in performance trade-offs that we attribute to selection for gliding in ground effect. Forewing shape differed between sexes for all taxa, and male wing shapes were aerodynamically more efficient for gliding flight than corresponding females. This suggests selection acts differentially on male and female wing shapes, reinforcing the idea that sex-specific flight behaviours contribute to the evolution of sexual dimorphism. Our study indicates that wing shapes in Haeterini butterflies evolved in response to habitat-specific flight behaviours, namely gliding in ground effect along the forest floor, resulting in ecomorphological partitions of taxa in morphospace. The convergent flight behaviour and wing morphology between tribes of Satyrinae suggest that the flight environment may offset phylogenetic constraints. Overall, this study provides a basis for exploring similar patterns of wing shape evolution in other taxa that glide in ground effect.

Diversification of Morpho butterflies (Lepidoptera, Nymphalidae): a re-evaluation of morphological characters and new insight from DNA sequence data

This study compiles previously published morphological, colour and behavioural characters and includes new DNA sequence data for eight markers (one mitochondrial and seven nuclear) to re‐evaluate phylogenetic relationships and estimate times of divergence for Morpho butterflies using parsimony and Bayesian methods. We note an effect of missing data on phylogenetic inference and calculations of Partitioned Bremer Support. Morphology and DNA trees were moderately congruent, and the combined analyses of all data included elements of both sources. Both morphology and DNA support the monophyly of Morpho and the early separation of the sister pair M. marcus plus M. eugenia, but trees from different data sources are congruent mostly at derived nodes, and differ at several internal nodes. The analyses of combined data indicate that Morpho is composed of four clades each of which include one or more previously proposed subgenera. The subgenera Pessonia and Morpho were not monophyletic, and to address this issue we propose that Pessonia, syn.nov. be subsumed within Morpho. The ancestor of Morpho probably arose during the Oligocene, and most diversification seems to have occurred during the late Miocene. S‐DIVA analysis suggests eastern Andean region as the ancestral area for Morpho, and that the South American Atlantic Forest was colonized multiple times.

Proboscis morphology and its relationship to feeding habits in noctuid moths.

Abstract This study describes proboscis morphology and identifies morphometric differences among five species of noctuid moths with different feeding habits (fruit versus nectar-feeding). Morphological and morphometric parameters were analyzed using scanning electron microscopy and light microscopy. Measurements included: galea height in ten sites from base to tip, total proboscis length, and length of the distal region that contains large sensilla styloconica and / or tearing hooks and erectible barbs. Both morphometric and morphological differences were identified among species within and between feeding guilds, and these results are discussed in light of the feeding habits of each species.

An illustrated key to male Actinote from Southeastern Brazil (Lepidoptera, Nymphalidae)

Em certas epocas do ano, as borboletas do genero Actinote podem ser muito abundantes no sudeste do Brasil, constituindo, entao, elementos conspicuous da nossa fauna. Sua coloracao alar permite a separacao em cinco padroes basicos, provavelmente relacionados com o mimetismo. Alem disso, series de exemplares coletados no campo ou criados em laboratorio demonstram claramente que existe uma grande variabilidade intra-especifica na coloracao alar destas borboletas. Em conjunto, a semelhanca entre as especies (mimetismo) e variacao intra-especifica dificultam o reconhecimento das especies de Actinote. Esta chave tem como objetivo auxiliar a identificacao dos machos destas borboletas e, para tanto, sao apresentados caracteres diagnosticos, ilustracoes das asas e genitalia de 22 especies. Esperamos que a clara identificacao destas especies possa revelar a presenca de especies ainda nao reconhecidas no sudeste do Brasil.

Both Palatable and Unpalatable Butterflies Use Bright Colors to Signal Difficulty of Capture to Predators

Birds are able to recognize and learn to avoid attacking unpalatable, chemically defended butterflies after unpleasant experiences with them. It has also been suggested that birds learn to avoid prey that are efficient at escaping. This, however, remains poorly documented. Here, we argue that butterflies may utilize a variety of escape tactics against insectivorous birds and review evidence that birds avoid attacking butterflies that are hard to catch. We suggest that signaling difficulty of capture to predators is a widespread phenomenon in butterflies, and this ability may not be limited to palatable butterflies. The possibility that both palatable and unpalatable species signal difficulty of capture has not been fully explored, but helps explain the existence of aposematic coloration and escape mimicry in butterflies lacking defensive chemicals. This possibility may also change the role that putative Müllerian and Batesian mimics play in a variety of classical mimicry rings, thus opening new perspectives in the evolution of mimicry in butterflies.

Molecular phylogeny and higher systematics of the metalmark butterflies (Lepidoptera: Riodinidae): Molecular phylogeny and systematics of Riodinidae

Riodinidae is a highly diverse butterfly family with the majority of its genera restricted to the Neotropics and, despite previous efforts, its higher systematics remains unresolved. Here, we propose a novel phylogenetic hypothesis, based on a comprehensive sample of riodinids, primarily from the Neotropics, covering 67% of all genera and all of the major lineages. We sequenced nine molecular markers and estimated resulting phylogenies with maximum likelihood and Bayesian approaches, using both timed trees and time‐independent trees. We based calibration on three fossil Riodinidae, and reassessed the position of the oldest fossil. We also incorporated 52 samples from a previous study providing a comprehensive maximum likelihood tree for 304 species comprising 80% of all genera. We propose a new higher classification of the Riodinidae with two subfamilies: the Nemeobiinae, including the Old World riodinids and their Neotropical sister Euselasia Hübner; and the Riodininae, comprising all remaining genera. We divided Riodininae into nine tribes (including four new tribes: Calydnini Seraphim, Freitas & Kaminski trib.n.; Sertaniini Seraphim, Freitas & Kaminski trib.n.; Dianesiini Seraphim, Freitas & Kaminski trib.n.; and Emesidini Seraphim, Freitas & Kaminski trib.n.), with Mesosemiini and Nymphidiini further subdivided into two and seven subtribes (including three new subtribes for Nymphidiini: Zabuellina Seraphim, Freitas & Kaminski subtrib.n.; Pachythonina Seraphim, Freitas & Kaminski subtrib.n.; and Pandemina Seraphim, Freitas & Kaminski subtrib.n.). Although our phylogenetic hypotheses are generally congruent with the analyses by Espeland et al. (2015), the comprehensive taxon sampling employed here constitutes a large step towards a stable tribal‐level classification. All taxonomic changes are summarized in a checklist. Despite most genera being restricted to tropical South and Central America, the oldest known fossil of Riodininae belongs to the Green River formation (42.6–50.2 Ma) in North America. Accordingly, we reassess the familys crown age at 56 Ma (52.4–60.7 Ma), which is at variance with previous dating using secondary calibrations and a different subset of genes. Nonmonophyletic riodinid genera are ubiquitous, and several groups need further revision, including groups revised recently. Our results point to the need for integrative taxonomy, as adult morphology seems to be have been exhausted as a single data source in this family.