The evolution of the laterophysic connection with a revised phylogeny and taxonomy of butterflyfishes (Teleostei: Chaetodontidae)

The higher‐level relationships of butterflyfishes were examined using 37 morphological characters. This analysis combines characters derived from a histological study describing variation in the morphology of the laterophysic connection (an association between the swim bladder and the lateral‐line canals) with previously described morphological characters. The phylogenetic analysis resulted in four equally parsimonious trees that only differed in the placement of two of the 11 chaetodontid genera (Amphichaetodon and Forcipiger). We compare our analysis with previous hypotheses, present a new taxonomy consistent with the proposed cladistic relationships, and diagnose Chaetodon with five unreversed synapomorphies, including the evolution of characters composing the laterophysic connection. A new character‐based diagnosis of Chaetodon is provided and species are allocated accordingly; Chaetodon now includes the former Parachaetodon ocellatus and excludes the former subgenera Prognathodes and Roa. The evolution of the laterophysic connection is examined by optimizing character‐state transformations on the new hypothesis of relationships.     

Phylogeny of the family Congiopodidae (Perciformes: Scorpaenoidea), with a proposal of new classification

The phylogenetic relationships of the family Congiopodidae are inferred based on morphological characters. The monophyly of this family is supported by 13 unambiguous apomorphic characters, including four autapomorphies among the superfamily Scorpaenoidea. The Congiopodidae shares 26 apomorphic characters with other scorpaenoid taxa, and these characters are considered to also support the monophyly of the family. Upon completion of the phylogenetic analysis using the characters in 39 transformation series, it was assumed that the family is unambiguously supported by five characters (and also by three and one characters when ACCTRAN and DELTRAN are used, respectively) and is branched into two major clades, including Congiopodus and Alertichthys plus Zanclorhynchus, respectively. Based on the phylogenetic relationships, a new classification, recognizing two subfamilies (Congiopodinae and Zanclorhynchinae) in the family Congiopodidae, is proposed. The genus Perryena, that was recently inferred being closely related to the Tetrarogidae (although many authors included it in the Congiopodidae), is provisionally placed into the Congiopodidae as incertae sedis.     

Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al. (1988): phylogenetic systematics and revised classification

Taxonomic schemes for the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al., (1988) have been unstable due to the large number of genera and the paucity of known reliable characters. Reliable characters are essential when using phylogenetic inference in developing a natural classification. Morphological and developmental studies using light, scanning and transmission electron microscopy have revealed the new characters of host response, en face patterns, phasmid structure and female cuticular layers. These techniques also gave us insight into the homoplasy and polarity of many characters, revealed previously undetected character states and clarified misinterpreted character states. A matrix with the 19 most reliable characters is proposed for 20 operational taxonomic units (OTUs) and we employ this matrix for comparing computer generated phylogenetic analyses of the PHYLIP and PAUP packages. PAUP was deemed the more reliable parsimony algorithm for phylogenetic analysis of the Heteroderinae (Fink, 1986; Platnick, 1987). Monophyly of Atalodera + Sherodera + Thecavermiculatus (tribe Ataloderini), and Cactodera + Heterodera + Afenestrata, as well as Punctodera + Globodera + Dolichodera is supported by both programs. Most importantly, analyses strongly support monophyly of all cyst-forming genera (tribe Heteroderini) contrary to previous hypotheses of repeated evolution of the cyst (Wouts, 1985). In addition, monophyly of the Heteroderini with the Ataloderini is demonstrated. PAUP indicates monophyly of Sarisodera + Rhizonema + Bellodera + Hylonema and Ekphymatodera (tribe Sarisoderini new rank). Monophyly of the Sarisoderini was at first only weakly supported, but, subsequently, the reduced width of the submedial lips of second stage juveniles and males was recognized as a synapomorphy which strengthened subsequent PAUP trees and monophyly of the tribe. The present study rejects as paraphyletic or polyphyletic several previously proposed combinations, including Thecavermiculatus sequoiae (versus Rhizonema sequoiae), Sarisodera africana (versus Afenestrata africana), Dolichodera andinus (versus Thecavermiculatus andinus). The question whether T. andinus is a distinct genus, was not resolved due to insufficient data. PAUP supports our previous observations that Cactodera betulae is intermediate in a transformation series between other Cactodera and Heterodera: it also indicates these species as bring monophyletic with Heterodera + Afenestrata, but not with other Cactodera. Although these phylogenetic analyses strongly support some relationships, they indicate unresolved alternative hypotheses for others. Meloidodera (tribe Meloidoderini) and Cryphodera (tribe Cryphoderini) must be investigated for consideration of a possible synapomorphy not included in the present data matrix. Future studies are proposed to more clearly define the monophyly of the Heteroderini, as well as the Sarisoderini. Tests are also proposed to clarify questions of the monophyly of Verutus (tribe Verutini new rank) with the Heteroderinae versus other Tylenchida.     

Redefinition and phylogenetic relationships of the family Pinguipedidae (Teleostei: Perciformes)

The phylogenetic relationships of the family Pinguipedidae plus the genus Cheimarrichthys von Haast, 1874, were studied to redefine the family. Based on a phylogenetic analysis using derived characters belonging to 21 transformation series, accepting the monophyly of Pinguipedidae plus Cheimarrichthys provisionally for the operational procedure of the analysis, it was hypothesized that Cheimarrichthys is not closely related to Parapercis Bleeker, 1863, although these genera had been considered as having a close affinity. Although the Pinguipedidae and Cheimarrichthys share nine derived characters, it was determined that these characters are also found in other trachinoid families. In addition, several families, such as the Leptoscopidae, Uranoscopidae, and Trichodontidae, have many more derived characters in common with Cheimarrichthys than the pinguipedids have with that genus. The conclusion drawn here is that it is not parsimonious to recognize the monophyly of the Pinguipedidae and Cheimarrichthys based on these nine derived characters, and that these characters are not compelling enough to link these groups. The redefined Pinguipedidae is proposed to include the following five genera: Parapercis, Prolatilus Gill, 1865, Pinguipes Cuvier, 1829, Pseudopercis de Miranda-Ribeiro, 1903, and Kochichthys Kamohara, 1960. Cheimarrichthys, excluded from the Pinguipedidae, is put into its own family, Cheimarrichthyidae.     

Mycoses in red snapper (Lutjanus campechanus) caused by two deuteromycete fungi (Penicillium corylophilum and Cladosporium sphaerospermum)*

We report two species of deuteromycete fungi (Penicillium corylophilum and Cladosporium sphaerospermum) concurrently infecting the swim bladder and posterior kidney and causing erratic behavior in two specimens of wild-caught, tank-held red snapper (Lutjanus campechanus). Lesions produced by both species infiltrated the immediately surrounding tissue and produced severe pathological changes; however, the infection apparently was not systemic. Only P. corylophilum grew in the initial culture from the swim bladder and only C. sphaerospermum grew in the initial culture from the kidney. Infection may have occurred upon penetration of a syringe to deflate the swim bladder. There was no horizontal transmission to 13 other specimens of red snapper held in the same tank. This suggests that these fungi are not primary pathogens. Injection of each species into various sites in the Gulf killifish, Fundulus grandis, failed to produce infections within 1 month, suggesting differences in susceptibility among species.     

Phylogeny of Callibaetis (Ephemeroptera: Baetidae) based on morphology of nymphs and adults

Callibaetis is considered to be one of the most problematic genera among mayflies by a series of taxonomic inaccuracies that have accumulated over the last two centuries. Despite these taxonomic problems, two independent hypotheses of species groups have been proposed. In the first hypothesis, three species groups for North America were proposed, and in the second, three species groups were proposed for South American. In these hypotheses, the generic delimitation and monophyly of Callibaetis have not been evaluated under a cladistic framework. Taking this into account, the objectives of this study were to verify the monophyly of Callibaetis and whether the groups of species proposed for the genus are corroborated as natural. The matrix included 128 morphological characters and one habitat character, 119 discrete characters were compiled (101 of nymphs and 18 of adults), and six imaginal characters are related to pigmentation pattern. Continuous characters were ratios and were represented by 10 characters (nine for nymphs and one for adults). The data set was analysed under implied weights. Group support was estimated with relative Bremer support and frequency differences. The results corroborated the monophyletic nature of Callibaetis and the generic status of Callibaetoides; however, the groups proposed for North and South American Callibaetis species were not corroborated. Our study indicated four groups of species for the genus, which we proposed as subgenera: Callibaetis, Abaetetuba subgen. n. Aiso subgen. n. and Cunhaporanga subgen. n.     

Cynanchum and the Cynanchinae (Apocynaceae – Asclepiadoideae): a molecular, anatomical and latex triterpenoid study

The phylogeny of the genus Cynanchum s. str. is studied using cpDNA spacers and ITS. Morphological, anatomical and latex triterpenoid data are interpreted in light of the molecular results, and discrepancies are discussed. Vegetative characters are better indicators of relationship than floral characters, especially corona characters. The monophyly of all Malagasy species and, nested within the latter, of all stem-succulent taxa is ascertained and the genera Folotsia, Karimbolea, Platykeleba and Sarcostemma are subsumed under Cynanchum. One African species, C. galgalense, is excluded from Cynanchum.     

Systematics of the tribe Trematocarini (Perciformes: Cichlidae) from Lake Tanganyika, Africa

 Phylogenetic relationships among eight Trematocara species and a single Telotrematocara species included in the Tanganyikan cichlid tribe Trematocarini were investigated on the basis of morphological features. The monophyly of the tribe is supported by the presence of hypertrophied sensory pores on the head, tendon “c” of adductor mandibulae section 1, a single scale row between the upper lateral line and body axis, great depth of the anteriormost infraorbital (reversed in Trematocara caparti and T. stigmaticum), and the absence of a lower lateral line. Trematocara is paraphyletic unless Telotrematocara is treated as a junior synonym. Received: December 10, 2001 / Revised: March 18, 2002 / Accepted: April 4, 2002     

Tiny steps towards greater knowledge: An osteological review with novel data on the Atlantic Forest toadlets of the Brachycephalus ephippium species group

Brachycephalus is a small, endemic genus of anurans that occur throughout the Brazilian Atlantic Forest. Recent analyses corroborated the monophyly of two species groups within this genus (B. ephippium and B. pernix), whereas the B. didactylus group appears to be polyphyletic. Herein, we compare and describe the skeletal system of all species from the Brachycephalus ephippium species group. We investigated diagnostic characters that are potentially useful to delimit similar species, confirmed the previously proposed genus definition and added six extra characters. We propose an osteological diagnosis for each species of the B. ephippium group, evaluating and describing intraspecific variations. Our results suggest that adults of B. ephippium group have ornamented parotic plates, ornamented post-orbital crests, absence of quadratojugal and neopalatines, distal end of the otic ramus of squamosal expanded towards the parotic plate, reduced zygomatic ramus of the squamosals, posterolateral projection of the crista parotica, presence of ornamented spinal plates (except B. alipioi) and ornamented paravertebral plates. We highlighted the importance of including more than one specimen per species when dealing with miniaturized taxa. A comparison with other Brachycephalus species groups and with Ischnocnema was also provided. Lastly, we suggest some characters to be included in future phylogenetic analysis.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

Reconciling taxonomy and phylogeny in the bristleworm family Eunicidae (polychaete,Annelida)

Eunicid annelids inhabit diverse marine habitats worldwide, have ecological and economic importance and have been pictured in the news as giant predator worms. They compose a traditional stable taxon recently supported as monophyletic but characterized by plesiomorphies. Most genera within the family have been recovered as paraphyletic in previous studies. We present a phylogenetic hypothesis for eunicid based on molecular (COI, 16S rDNA, 18S rDNA) and morphological data (213 characters), including an explicit attempt to account for serial homology. Eunicidae as well as monophyletic genera Marphysa sensu stricto and Lysidice is redefined based on synapomorphies. Nematonereis is synonymized to Lysidice. Leodice and Nicidion are resurrected to name monophyletic groups including species previously included in Eunice and Marphysa sensu lato. Traditional diagnostic characters such as the absence/presence of peristomial cirri, lateral antennae and branchiae are homoplasies and not informative at the generic level. Different coding of traditional characters (i.e. articulation of prostomial appendages) and novel characters of prostomial features and regionalization of the body support the monophyly of the family and genera level clades. Thus, the phylogenetic hypothesis presented here and the evolution of characters provided background information for taxonomic changes yielding evolutionary meaningful classification and diagnoses for the family and genera.     

A phylogenetic study of the Thygater‐Trichocerapis group and new scopes for the subgenera of Thygater Holmberg, (Hymenoptera,Apidae)

The bee tribe Eucerini is a large monophyletic taxon occurring in all continents, except Oceania and Antarctica, but far more diverse in the Americas and, especially, in the Neotropics. The phylogenetic relationships within its subtribe Eucerina, which contains the bulk of Eucerini diversity, is poorly understood, this being especially true for the relationships among its Neotropical representatives, which have been poorly represented in all phylogenetic studies, including Eucerini, to date. This leads to a generalized lack of confidence on the monophyly of the groups currently accepted as genera and subgenera. Here, a phylogenetic study based on three molecular markers (COI, 18S, and 28S, totalling about 1700 bp) and 58 morphological characters is presented as a contribution to the understanding of the relationships of the so‐called Thygater‐Trichocerapis group, and especially of the genus Thygater Holmberg, which has not previously been extensively sampled. Representatives of Trichocerapis, including its monotypic subgenus T. (Dithygater), are included for the first time in a phylogenetic study. The main results were: (i) support for the monophyly of the Thygater‐Trichocerapis group; (ii) support for the monophyly of Thygater; (iii) recognition of two main clades in Thygater, each one containing the type species of one of the previously recognized subgenera; and (iv) additional support for the position of Alloscirtetica as sister to all remaining Eucerina. Based on these results a redefinition of the scope of the two subgenera of Thygater is proposed, with changes in the subordination of three of its species, T. (Nectarodiaeta) chaetaspis comb.n. , T. (Nectarodiaeta) paranaensis comb.n. and T. (Thygater) mexicana comb.n.     

Evolution of Philopotinae,with a revision and phylogeny of the New World spider fly genus Philopota Wiedemann (Diptera,Acroceridae)

Philopotinae are hunchbacked spider flies represented by 63 fossil and extant species in 15 genera worldwide. Philopota Wiedemann, 1830, is the most species‐rich genus within the subfamily. Here, the evolution of Philopotinae is discussed, and a revision and phylogeny of Philopota based on adult morphology are presented. Nine of the 12 extant Philopotinae genera were included in our analysis, and 22 species were recognized in Philopota, of which 13 are described as new. Seven new synonymies are proposed. The phylogenetic analysis included 33 terminal taxa (22 ingroup and 11 outgroup species) and used 53 morphological characters, resulting in a single most parsimonious tree under equal weights. The monophyly of Philopota is recovered, and the Palaearctic genus Oligoneura is hypothesized as its sister‐group.     

Intra- and inter-specific variation in buoyancy of some estuarine fishes

Synopsis Buoyancy was measured on eight species of estuarine fishes that were caught in 1 m depth or less. Mean buoyancies of the physoclists Fundulus heteroclitus, F. majalis, Cyprinodon variegatus and Leiostomus xanthurus were similar and ranged from –6.5 to –18.0 kiloPascals below atmospheric pressure at sea level. Menidia menidia and Pomatomus saltatrix measured –36.6 and –46.1 kPa, respectively. Two physostomes, Brevoortia tyrannus and Anchoa mitchilli, measured + 2.9 and –23.5 kPa, respectively, but the latter probably releases air when handled.The four most buoyant physoclist species live near the bottom in areas that receive daily tide induced currents. Negative buoyancy probably functions in them as in stream dwelling minnows and salmonids, which respond to currents by decreasing their buoyancy. The pronounced negative buoyancy of M. menidia may be a response to a preference for habitat where the currents are stronger, P. saltatrix, which can secrete gas into the swim bladder at the fastest rate known for any fish, combines high secretion (and resorption) rates with marked negative buoyancy. This enables it to quickly change depths over a wide vertical range, without overexpanding the swim bladder to cause positive buoyancy.     

The Phylogeny of the Myrmecophagidae (Mammalia, Xenarthra, Vermilingua) and the Relationship of Eurotamandua to the Vermilingua

A cladistic investigation of the phylogenetic relationships among the three extant anteater genera and the three undoubted extinct myrmecophagid genera is performed based upon osteological characteristics of the skull and postcranial skeleton. One hundred seven discrete morphological characters are analyzed using the computer program PAUP. Characters are polarized via comparison to the successive xenarthran outgroups Tardigrada (represented by the living sloth Bradypus) and Cingulata (represented by the recent armadillos Dasypus and Euphractus). The analysis results in a single most-parsimonious tree (TL = 190, CI = 0.699, RI = 0.713). The tree corroborates the monophyly of the subfamilies Cyclopinae and Myrmecophaginae, the former including the extant Cyclopes and the Pliocene genus Palaeomyrmidon. Within the Myrmecophaginae the Miocene genus Protamandua is the sister taxon to a clade including the remaining three genera. The recent Tamandua is in turn the sister taxon to the extant Myrmecophaga plus the Pliocene genus Neotamandua. Contrary to the suggestions of recent authors, weak support is provided for the taxonomic distinctiveness of the latter genus from the recent Myrmecophaga. The monophyly of the Myrmecophagidae is supported by 15 unequivocal synapomorphies. The monophyly of the Cyclopinae and Myrmecophaginae is supported by 3 and 13 unambiguous synapomorphies, respectively. The enigmatic Eocene genus Eurotamandua, from the Messel fauna of Germany, is coded for the 107 morphological characters above and included in two subsequent PAUP analyses. The palaeanodont Metacheiromys is also added to these two analyses as a nonxenarthran outgroup to test for the possibility that Eurotamandua lies outside the Xenarthra. In the first analysis, Eurotamandua is constrained a priori to membership in the Vermilingua. The single most-parsimonious tree (TL = 224, CI = 0.618) that results places Eurotamandua as the sister group to the remaining anteater genera, contra Storch and Habersetzer's (1991) assignment of Eurotamandua to the vermilinguan subfamily Myrmecophaginae. Eurotamandua shares six unequivocal synapomorphies with other anteaters, including the absence of teeth and the presence of a lateral tuberosity on the fifth metatarsal. The remaining vermilinguans are united by 11 unequivocal synapomorphies, plus an additional 10 ambiguous synapomorphies. In the second analysis, the position of Eurotamandua is unconstrained. The resulting single most-parsimonious tree (TL = 219, CI = 0.632) places Eurotamandua outside Vermilingua as the sister group to the Pilosa (Vermilingua plus Bradypus). The monophyly of this node is supported by four unambiguous synapomorphies in the unconstrained analysis. Further manipulation of this second analysis shows that placement of Eurotamandua as the sister group to the Xenarthra or to the Palaeanodonta adds three steps to the shortest tree but is more parsimonious than its placement as a sister group to the Vermilingua is the previous analysis. The addition of pangolins to the analysis does little to alter the major phylogenetic conclusions of the study. The allocation of Eurotamandua to the Xenarthra, but as a sister group to the Pilosa, is a novel arrangement which leaves open the biogeographic question of how a xenarthran reached Western Europe during the Eocene.     

An assessment of the status of Polycirridae genera (Annelida: Terebelliformia) and evolutionary transformation series of characters within the family

A phylogenetic analysis was performed to determine the monophyly of non‐monotypic genera of the terebelliform family Polycirridae, i.e. Polycirrus, Amaeana, Lysilla, and Hauchiella, and the evolution of characters among members of this clade. The monotypic genera, Enoplobranchus and Biremis, were also included, together with members of both known species in Hauchiella. Representative species were included for remaining genera: 14 species of Polycirrus, six species of Amaeana, and six species of Lysilla. Out‐groups consisted of representatives of Spionidae, Cirratulidae, and Sabellariidae, as well as several species of Telothelepodidae. A total of 40 in‐ and out‐group species were coded for 50 subjects (‘characters’) and 117 subject–predicate relationships (‘states’). Although results are consistent with recent phylogenetic studies within Terebelliformia that suggest Polycirridae monophyly, only Hauchiella was found to be monophyletic, albeit part of the more inclusive clade comprising remaining polycirrid genera. Evolutionary transformation series are discussed for selected characters in relation to the non‐monophyly of Polycirrus, Lysilla, and Amaeana. Implications for the use of supraspecific taxa as ‘taxonomic surrogates’ are highlighted. The definition of Polycirridae is emended. © 2015 The Linnean Society of London     

Phylogenetics and biogeography of the broad-nosed bats, genus Platyrrhinus (Chiroptera: Phyllostomidae)

The Neotropical broad-nosed bats, genus Platyrrhinus, represent a well-defined monophyletic group of 14 recognized species. A recent study of morphological characters confirmed Platyrrhinus monophyly and species diagnosis, but offered little support to their intra-specific relationships. We conducted phylogenetic analyses of the genus, using dense taxonomic sampling in combination with four gene sequences representing both mitochondrial and nuclear DNA transmission systems. Our aim was to elucidate the phylogenetic structure among species, using the resulting 3341 bp of DNA. Maximum parsimony, maximum likelihood, and Bayesian inference analyses produced similar topologies that confirm the monophyly of the genus Platyrrhinus and strongly support many previously unrecognized groups. Paraphyly of Platyrrhinus helleri and the unclear position of P. brachycephalus in the clades were also apparent in the data. Our biogeographical analysis suggests a Brazilian Shield origin for Platyrrhinus, followed by subsequent radiations of lineages in the Amazon Basin and Andes. Secondary dispersal from Amazonian and Andean centers is responsible for the Platyrrhinus inhabiting the Guianan Shield and the Pacific lowlands and Central America, respectively.     

Artificial neural networks for fly identification: A case study from the genera <Emphasis Type="Italic">Tachina</Emphasis> and <Emphasis Type="Italic">Ectophasia</Emphasis> (Diptera,Tachinidae)

The classification methodology based on morphometric data and supervised artificial neural networks (ANN) was tested on five fly species of the parasitoid genera Tachina and Ectophasia (Diptera, Tachinidae). Objects were initially photographed, then digitalized; consequently the picture was scaled and measured by means of an image analyser. The 16 variables used for classification included length of different wing veins or their parts and width of antennal segments. The sex was found to have some influence on the data and was included in the study as another input variable. Better and reliable classification was obtained when data from both the right and left wings were entered, the data from one wing were however found to be sufficient. The prediction success (correct identification of unknown test samples) varied from 88 to 100% throughout the study depending especially on the number of specimens in the training set. Classification of the studied Diptera species using ANN is possible assuming a sufficiently high number (tens) of specimens of each species is available for the ANN training. The methodology proposed is quite general and can be applied for all biological objects where it is possible to define adequate diagnostic characters and create the appropriate database.     

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.