Keith Willmott

University of Florida | UF · Department of Natural History

Understanding the mechanisms underlying species distributions and coexistence is both a priority and a challenge for biodiversity hotspots such as the Neotropics. Here, we highlight that Müllerian mimicry, where defended prey species display similar warning signals, is key to the maintenance of biodiversity in the c. 400 species of the Neotropical...

Species are the fundamental units of life and evolution. Their recognition is essential for science and society. Molecular methods have been increasingly used for the identification of animal species, despite several challenges. Here, we explore with genomic data from nine animal lineages a set of nuclear markers, namely metazoan-level universal si...

The richest butterfly communities in the world are found in the Amazon rainforest. Despite of this, and the importance of species inventories for the knowledge of diversity patterns, there are few comprehensive lists of butterflies for localities in the Brazilian Amazon. Here, we present an updated list of the butterflies of Cristalino Lodge (Alta...

A comparative analysis of high-Andean Pierina was carried out, including a total of 25 species. Based on morphological evidence, with an emphasis on venation and genitalia and molecular data, using three genetic markers, we confirm the recent subjective synonymy of the generic names Tatochila Butler, 1870, Piercolias, Staudinger, 1894, Hypsochila U...

Temperature is thought to be a key variable explaining global patterns of species richness. However, to investigate this relationship carefully, it is necessary to study clades with broad geographic ranges that are comprised of species inhabiting diverse biomes with well-characterized species ranges. In the present study, we investigate the link be...

The species-level classification of the "celia clade" of the nymphalid butterfly genus Pseudodebis Forster, 1964, is revised as part of ongoing revisionary work on this genus. The "celia clade" contains three species, of which two, Pseudodebis dar-renthroopi Nakahara & Willmott, n. sp. and P. tigrillo Nakahara & Willmott, n. sp., are described and...

A new ithomiine butterfly, Mechanitis lysimnia tapajona Freitas & Mota ssp. nov., is described from four localities in southern Amazonia. This taxon is very similar to the subspecies Mechanitis lysimnia nesaea, but can be distinguished by the orange hindwing discal cell (which is yellow in M. l. nesaea). The recognition of Mechanitis lysimnia tapaj...

Insects are facing severe threats of extinction. Long-term data needed to develop effective conservation strategies are not yet available for precisely those areas where biodiversity peaks and negative impacts on species are particularly strong, such as Neotropical forests. One strategy to develop long-term monitoring programmes worldwide has focus...

We here establish a new genus in the nymphalid butterfly subtribe Euptychiina, Cisandina Nakahara & Espeland, n. gen. to harbor five species hitherto placed within two polyphyletic genera, namely Magneuptychia Forster, 1964 and Euptychoides Forster, 1964. We compiled data from over 350 specimens in 17 public and private collections, as well as DNA...

We here describe a new satyrine species in the genus Chloreuptychia Forster, 1964, C. gordonsmalli Thurman, MacDonald & Nakahara, n. sp., based on four historical specimens all originating from central Panama. Despite the lack of genetic data for this new taxon and the polyphyletic nature of Chloreuptychia, some genitalic characters suggest that th...

Aim The biodiversity crisis has highlighted the need to assess and map biodiversity in order to prioritize conservation efforts. Clearwing butterflies (tribe Ithomiini) have been proposed as biological indicators for habitat quality in Neotropical forests, which contain the world's richest biological communities. Here, we provide maps of different...

The Huancabamba Deflection in the Andes of northern Peru and southern Ecuador is a pivotal area for Neotropical biogeography, where exceptional biodiversity coincides with high rates of endemism. These characteristics are well illustrated within the montane butterfly genus Manerebia Staudinger (Nymphalidae, Satyrinae). Here, six new, apparently end...

A new satyrine butterfly species, Splendeuptychia tupinamba Freitas, Huertas & Rosa, sp. nov. (Nymphalidae: Satyrinae), is described. This species is found throughout a large geographical range in Brazil, Argentina and Paraguay, predominantly in the Cerrado domain, with some records in neighboring Amazonia and Atlantic Forest. Morphology and molecu...

The Heliconiini genera Agraulis and Dryas are widely distributed throughout the Neotropics and into adjacent temperate regions, and although they are currently treated as monotypic, both show significant geographic phenotypic variation. In this work, we employ six genetic markers (4199 bp), two mitochondrial and four nuclear, to perform coalescent...

A new species of Taydebis Freitas, 2003 from south Brazil is described using comparative morphology and species distributions. Also, based on morphology, we transfer Neonympha melobosis Capronnier, 1874 (formerly placed in Paryphthimoides Forster, 1964) to Taydebis, and recognize Euptychia peculiaris Butler, 1874 as its junior synonym (syn. nov.)....

A new subspecies of Moschoneura pinthous (Linnaeus, 1758), M. p. carmenae Neild & Losada, ssp. nov., is described from the Orinoco Delta in northeastern Venezuela. The adults, male genitalia, and DNA barcode sequences are documented and compared with neighboring M. p. pinthous from southern Venezuela and the Guianas, and with M. pinthous subspecies...

Species are the fundamental units of life and their recognition is essential for science and society. DNA barcoding, the use of a single and often mitochondrial gene, has been increasingly employed as a universal approach for the identification of animal species. However, this approach faces several challenges. Here, we demonstrate with empirical d...

We name and describe a new nymphalid butterfly species, Pseudodebis hartmanni Nakahara & MacDonald, n. sp. (Satyrinae: Satyrini) based on differences in its wing pattern and DNA from the closely related species. Phylogenetic analysis suggests that this Central American species belongs to a strongly supported clade that also includes two sympatric s...

Most research on aposematism has focused on chemically defended prey, but the signalling difficulty of capture remains poorly explored. Similar to classical Batesian and Müllerian mimicry related to distastefulness, such ‘evasive aposematism' may also lead to convergence in warning colours, known as evasive mimicry. A prime candidate group for evas...

DNA ‘barcoding’ has contributed greatly to resolving species limits in rapidly diverging tropical insect groups. Here, we explored species diversity in the widespread, cryptic Neotropical butterfly genus Hermeuptychia by generating 601 new sequences of the cytochrome c oxidase I (COI) barcode, tripling available information. Our dataset focused in...

The field of systematics and our understanding of phylogenetic relationships have been invigorated by the use of molecular data, but analyses based on DNA sequence data are not always corroborated by diagnostic morphological characters. In particular, several taxonomic changes in butterflies (Papilionoidea) have been made solely on the basis of mol...

We here propose a new monotypic butterfly genus Scriptor Nakahara & Espeland, n. gen. to accommodate a new species, S. sphenophorus Lamas & Nakahara, n. sp., described and named herein. Scriptor sphenophorus n. gen. and n. sp. is a relatively common and widespread butterfly species which is recovered as a member of the so-called ''Splendeuptychia c...

We here transfer an euptychiine taxon hitherto placed in the polyphyletic genus Magneuptychia Forster, 1964, to Caeruleuptychia Forster, 1964. Caeruleuptychia francisca (Butler, 1870), n. comb. is reclassified based on a morphology-based maximum likelihood analysis, which is consistent with ongoing analyses of molecular data. Two putative synapomor...

Vareuptychia Forster, 1964 stat. rest. is revalidated and comprises two species, V. similis (Butler, 1867) comb. rest. and V. themis (Butler, 1867) comb. nov. Vanima Zacca, Casagrande Mielke gen. nov. is described to contain Euptychia labe Butler, 1870 (the type species), E. palladia Butler, 1867 and E. lesbia Staudinger, [1886]. The taxonomy of th...

A new genus of Euptychiina, Emeryus Zacca, Casagrande & Mielke, gen. nov., is herein described using an integrative taxonomic approach to accommodate three species previously placed in Paryphthimoides Forster, 1964: Emeryus argulus argulus (Godart, [1824]), comb. nov. (type species), Emeryus difficilis (Forster, 1964), comb. nov. and Emeryus numeri...

Most research on aposematism has focused on chemically defended prey, but signalling difficulty of capture remains poorly explored. Similarly to classical Batesian and Müllerian mimicry related to distastefulness, such "evasive aposematism" may also lead to convergence in warning colours, known as evasive mimicry. A prime candidate group for evasiv...

We describe for the first time the immature stages of the Ecuadorian endemic urban butterfly, Catasticta flisa duna Eitschberger & Racheli, from Quito, Ecuador. A mistletoe species, Phoradendron nervosum Oliv. (Viscaceae), is identified as the host plant. Phoradendron nervosum is an aerial-stem hemiparasite in the order Santalales that is parasitic...

The latitudinal diversity gradient (LDG) is arguably one of the most striking patterns in nature. The global increase in species richness toward the tropics across continents and taxonomic groups stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluated several of these hypotheses to explain sp...

A systematic revision of Paryphthimoides , a satyrine genus widely distributed in Central and South America, is presented based on an integrative approach. Molecular phylogeny inferred using multiple methods with nuclear (GAPDH, RpS5) and mitochondrial (COI) genes indicates that Paryphthimoides , as currently circumscribed, is polyphyletic. A new t...

The male genitalic characters of Hexapoda are well known for their great taxonomic and systematic value. Despite insect male genitalia displaying large diversity, variation, and modification across orders, some structures are consistently present, and such characters can serve as the basis for discussion regarding homology. In the order Lepidoptera...

We provide the first comprehensive taxonomic revision of the poorly known South American butterfly genus Zischkaia Forster, 1964, hitherto regarded as including three described species. A phylogenetic analysis based on DNA sequence data shows that Zischkaia is monophyletic and consists of two morphologically diagnosable clades. Morphological charac...

Adelpha messana (C. Felder & R. Felder, 1867) is an uncommon forest species hitherto known from Central America to the Amazon basin. Here we report the species for the first time from the Atlantic Forest region of southeastern Brazil, where it occurs as a distinct subspecies which we describe and name as Adelpha messana seminivea Freitas, Willmott...

The landscape of the Neotropical region has undergone dynamic evolution throughout the Miocene, with the extensive Pebas wetland occupying western Amazonia between 23 and c. 10 Ma and the continuous uplift of the Andes mountains. The complex interaction between the Andes and Amazonia probably influenced the trajectory of Neotropical biodiversity, b...

Our goal was to test the hypothesis that assembly processes that limit species similarity (i.e., competition) predominantly occur in more "stable" abiotic environments, whereas habitat filtering (i.e., habitat characteristics) is a major driver of community composition within more variable environments at regional (e.g., aseasonal vs seasonal fores...

We analyzed the dynamics of multi-species butterfly communities along a climatic gradient with varying precipitation regimes for three consecutive years, and determine how climatic variables associate with observed butterfly seasonality. To provide a baseline for future studies of how climate change might affect these butterfly populations, we addi...

A new genus, Optimandes Marín, Nakahara & Willmott, n. gen., is described to contain the type species Neonympha eugenia C. Felder & R. Felder, 1867, its junior subjective synonym Euptychia phineus Butler, 1867, and its subspecies Euptychia transversa Weymer, 1911, which are transferred from the genus Euptychoides Forster, 1964. The sister species t...

We here propose a new, monotypic genus, Amiga Nakahara, Willmott & Espeland, gen. n., to harbor a common Neotropical butterfly, described as Papilio arnaca Fabricius, 1776, and hitherto placed in the genus Chloreuptychia Forster, 1964. Recent and ongoing molecular phylogenetic research has shown Chloreuptychia to be polyphyletic, with C. arnaca pro...

Relationships within satyrine butterflies have been notoriously difficult to resolve using both morphology and Sanger sequencing methods, and this is particularly true for the mainly Neotropical subtribe Euptychiina, which contains about 400 described species. Known larvae of Euptychiina feed on grasses and sedges, with the exception of the genus E...

Aim: Geographic distributions are driven by a combination of species sensitivity and exposure to climate. We quantified colour lightness, a trait that mediates the interaction between sensitivity and exposure, of diverse butterfly communities to test whether colour lightness is associated with community assembly across climate‐elevation gradients....

The evolution of reproductive isolation via a switch in mimetic wing coloration has become the paradigm for speciation in aposematic Heliconius butterflies. Here, we provide a counterexample to this, by documenting two cryptic species within the taxon formerly considered Heliconius demeter Staudinger, 1897. Amplified fragment length polymorphisms i...

The leafwing butterflies of the genus Zaretis Hübner, [1819] are revised based on morphological and molecular data. Species were delimited using morphology and 396 ‘DNA barcode’ sequences. Intraspecific and interspecific genetic distances, along with neighbor-joining, maximum likelihood and Bayesian inference analyses, were used to infer putative s...

Seven new euptychiine (Lepidoptera: Nymphalidae: Satyrinae) taxa are described and named herein, namely Harjesia argentata Nakahara, Zacca and Lamas, n. sp., Orotaygetis Nakahara and Zacca, n. gen., O. surui Nakahara, Zacca and Lamas, n. sp., Euptychoides sanmarcos Nakahara and Lamas, n. sp., Pseudeuptychia cuzquenya Nakahara and Lamas, n. sp., P....

The genus Cissia Doubleday, 1848 is revised and its monophyly is tested using Maximum Likelihood analysis based on three genes (COI, GAPDH and RpS5). A new taxonomic arrangement is proposed based on molecular and morphological evidence, and Cissia now comprises six species, C. penelope (Fabricius, 1775), C. pompilia (C. Felder & R. Felder, 1867), C...

A new species of Euptychiina, Erichthodes eremita Lamas, Willmott & Radford, n. sp., is described and illustrated. DNA sequence data suggest that the new species is sister to a species currently placed in Erichthodes Forster, 1964, although ongoing revision of the generic taxonomy of the subtribe might result in the reclassification of both of thes...

The discovery of a new species of satyrine butterfly, Magneuptychia louisammour Benmesbah & Zacca, sp. n., that is phenotypically similar to a sympatric species widely known as Magneuptychia ocypete (Fabricius, 1776), led to a review of the taxonomic status of M. ocypete. A neotype for Papilio ocypete Fabricius, 1776 is designated and its redescrip...

Despite more than a century of biological research on the evolution and maintenance of mimetic signals, the relative frequencies of models and mimics necessary to establish and maintain Batesian mimicry in natural populations remain understudied. Here we investigate the frequency-dependent dynamics of imperfect Batesian mimicry, using predation exp...

The Neotropical region is the most biodiverse on Earth, in a large part due to the highly diverse tropical Andean biota. The Andes are a potentially important driver of diversification within the mountains and for neighboring regions. We compared the role of the Andes in diversification among three subtribes of Ithomiini butterflies endemic to the...

Euptychia saltuensis Hayward, 1962, new synonym, currently regarded as a nomen dubium and possibly a junior subjective synonym of Yphthimoides manasses (C. Felder & R. Felder, 1867), is here treated as a junior subjective synonym of Yphthimoides patricia (Hayward, 1957), based on morphological characters of the male genitalia and the DNA barcode. T...