The generic classification of the Trochilini (Aves: Trochilidae): Reconciling taxonomy with phylogeny

Abstract

The generic nomenclature of the hummingbirds is unusually complicated. McGuire et al.’s (2014) recent phylogeny of the Trochilidae based on DNA sequence data has greatly clarified relationships within the family but conflicts strongly with the traditional classification of the family at the genus level, especially that of the largest and most recently derived clade, the Trochilini or “emeralds”. We recently presented a historical review of this classification and the generic modifications required by the Code of the International Commission on Zoological Nomenclature. Herein we present a revised generic classification of the Trochilini based upon McGuire et al.’s genetic data, while producing diagnosable generic groupings and preserving nomenclatural stability insofar as possible. However, this generic rearrangement has necessitated the resurrection of nine generic names currently considered synonyms, the synonymization of seven currently recognized genera and the creation of one new genus. The generic changes we recommend to the classification are drastic, and we summarize these in tabular form in comparison with the three most recent classifications of the Trochilini. Where appropriate, we outline alternatives to our proposed arrangement. The classification treats 110 species in 35 genera, including two species that remain unplaced for lack of genetic samples.

Full text

Accepted by P. Rasmussen: 16 Oct. 2017; published: 24 Nov. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2017 Magnolia Press
Zootaxa 4353 (3): 401
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424
http://www.mapress.com/j/zt/
Article
401
https://doi.org/10.11646/zootaxa.4353.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:54F45732-C58B-4FF5-8697-FC3F942445BC
The generic classification of the Trochilini (Aves: Trochilidae):
Reconciling taxonomy with phylogeny
F. GARY STILES
1
, J. V. REMSEN, JR.
2
& JIMMY A. MCGUIRE
3
1
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia. E-mail: fgstilesh@unal.edu.co
2
Museum of Natural Science & Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA.
E-mail: najames@LSU.edu
3
Museum of Vertebrate Zoology, University of California, Berkeley, California 94720, USA
Abstract
The generic nomenclature of the hummingbirds is unusually complicated. McGuire et al.’s (2014) recent phylogeny of the
Trochilidae based on DNA sequence data has greatly clarified relationships within the family but conflicts strongly with
the traditional classification of the family at the genus level, especially that of the largest and most recently derived clade,
the Trochilini or “emeralds”. We recently presented a historical review of this classification and the generic modifications
required by the Code of the International Commission on Zoological Nomenclature. Herein we present a revised generic
classification of the Trochilini based upon McGuire et al.’s genetic data, while producing diagnosable generic groupings
and preserving nomenclatural stability insofar as possible. However, this generic rearrangement has necessitated the res-
urrection of nine generic names currently considered synonyms, the synonymization of seven currently recognized genera
and the creation of one new genus. The generic changes we recommend to the classification are drastic, and we summarize
these in tabular form in comparison with the three most recent classifications of the Trochilini. Where appropriate, we
outline alternatives to our proposed arrangement. The classification treats 110 species in 35 genera, including two species
that remain unplaced for lack of genetic samples.
Key words: biogeography, classification, genetic relationships, hummingbirds, morphology, nomenclature
Introduction
A recent phylogeny based on DNA sequence data that included 275 species of hummingbirds (McGuire et al.
2014) revealed widespread polyphyly among many of the currently recognized genera in the Trochilidae. Nowhere
is the incongruence between the traditional classification and the phylogeny more acute than in the tribe Trochilini
(sensu McGuire et al. 2009; Dickinson & Remsen 2013), popularly called the “emeralds”, which constitute the
largest (over 100 species) clade in the Trochilidae. A thorough overhaul of the arrangement and circumscription of
the generic classification of the Trochilini is therefore urgently needed.
As a necessary first step, we reviewed the tangled history of the causes and extent of the problems with
hummingbird genera in general, with particular emphasis on the Trochilini (Stiles et al. 2017). Therein we noted
that as a whole, members of this tribe show far greater diversity in the colors and patterns of their plumages than in
their external morphology, perhaps reflecting their relatively restricted ecological and geographical distributions:
no member of the clade occupies the higher elevations in the tropics, nor the higher latitudes in the north or south.
Because previous classifications of the emeralds were largely based on plumage, and because the degree of
homoplasy in colors and patterns had been strongly underestimated, the resulting degree of correspondence of
these classifications with the genetic data is poor. The objective of this paper is to bring the classification of the
emeralds into as close an accord with the phylogeny as possible.

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Methods
Our basic premise in this revision is that the molecular phylogenetic tree of McGuire et al. (2014) represents the
best available estimate of phylogenetic relationships within the Trochilini. The branching pattern of this tree is
therefore our starting point, because it reveals the pattern of diversification within the family, shows most clearly
the homoplasy in many traditional characters previously used to define genera, and provides the rationale for many
new generic groupings. The resulting generic rearrangement has necessitated resurrecting several generic names
considered synonyms in current classifications; we have paid special attention to the type species previously
designated for these names. A companion paper (Stiles et al. 2017) provides the historical background for these
rearrangements through application of the International Commission on Zoological Nomenclature ICZN (1999)
Code, in the broader context of the development of hummingbird classifications in general. We emphasize that our
goals of producing morphologically and biogeographically diagnosable genera while maintaining stability in
generic nomenclature are to a considerable extent incompatible. This has obliged us to adopt a more flexible
treatment of branch lengths to avoid excessively diverse and undiagnosable genera or a multiplicity of small
genera, especially when this would require creating new generic names. Only in one case has creating a new genus
been necessary because neither of the two included species had been designated the type species of an applicable
generic name. For the new groupings required by the genetic tree, we have sought previously unappreciated
similarities among the included species to facilitate their diagnoses. When two or more possibilities exist for
defining generic circumscriptions, we have presented these as alternatives but have usually recommended that
which best preserves nomenclatural stability. Genetic data for several species in the Trochilini are still lacking, but
in nearly all cases inclusion of these species is consistent in morphology or other features with existing or new
generic groupings. Only two Middle American species of Trochilini without genetic samples remain unplaced
(incertae sedis) because we are reluctant to rely solely upon plumage characters subject to homoplasy for their
placement.
Here, we examine the current generic classification of the emeralds as presented by Peters (1945),
Schuchmann (1999) and Dickinson & Remsen (2013) with respect to the phylogeny estimated by DNA sequence
data (McGuire et al. 2014), and we recommend adjustments to the classification to bring it into accord with that
phylogeny. We emphasize that in general, genetic differences among many groups of emeralds contrast with their
limited morphological diversity, and so have preferred to recognize multiple genera in such cases to emphasize
ages of lineages. By contrast, several groups recognized as monophyletic in the phylogeny exhibit marked diversity
in plumage and shape, which therefore cannot be relied upon to delimit genera in such cases. In Appendix 1 we
give the original descriptions of all genera and of all type species in abbreviated form, but we present the full
citations in the Literature Cited section of the text below.
The most difficult aspect of this study has involved the consultation of the original references for the generic
circumscriptions and type species designations. For this, we found the digitalized versions of most of these through
the invaluable collections of the Biodiversity Heritage Library, Gallica and the Zoonomen index; for the most
complicated cases, including those of Lesson and Gould (who published their works in numerous installments for
which assigning dates was often difficult) we have relied on the reviews by Coues (1879), Cretella (2005) and
Dickinson et al. (2011).
Changes in generic names in the Trochilini mandated by the phylogeny
The genetic data of McGuire et al. (2014) reveals four major groups in the Trochilini, which for convenience we
will label A, B, C and D (Fig.1). The first three of these present relatively few problems in terms of classification,
but group D is by far the largest and most in need of a thorough revision of the current generic classification. For
each group below, we realign generic boundaries as required by the genetic data and discuss alternatives for
especially problematic cases.
Group A includes the currently recognized genera Cynanthus Swainson, 1827a, Cyanophaia Reichenbach,
1854, and Chlorostilbon Gould, 1853. A character shared by almost all genera in this group, with the probable
exception of Cyanophaia (see below) is a distinctive type of tomial serration (class A of Stiles & Rico, submitted).
This serration type has also evolved independently in all of clade 2 (mangos), twice in the hermits (Ramphodon and

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A NEW CLASSIFICATION OF THE TROCHI LINI
Glaucis), twice in the coquettes (Aglaiocercus and Sappho), and again twice independently in the Trochilini
(groups A and C below). One possible taxonomic treatment of Group A would be to expand Cynanthus to include
Chlorostilbon and Cyanophaia. Based on comparative branch lengths, however, a broadly defined Cynanthus
would mask considerable genetic diversity, thus favoring a classification that recognizes this diversity with
multiple genera.
In group A (Fig. 1), Cynanthus sordidus Gould is separated on a long branch, forming a separate subgroup A1,
sister to the rest of the group. This species was formerly (Ridgway 1911; Cory 1918) considered to represent a
monotypic genus Phaeoptila Gould, 1861, which was merged into Cynanthus without comment by Peters (1945).
We recommend returning it to its former status as a monotypic genus, as Phaeoptila sordida Gould, 1859. Its dull,
sexually monomorphic plumage is unique in the group, and it also has exceptionally well-developed tomial
serrations.
Subgroup A2 includes the Greater Antillean species of Chlorostilbon plus Cyanophaia Reichenbach, 1854,
endemic to the Lesser Antilles. Because the other species of Chlorostilbon (including its type species, mellisugus
Linnaeus, 1758, as determined by Zimmer 1950) are included in subgroup A4, another generic name is required for
the Antillean species; the candidates are Riccordia Reichenbach,1854, Sporadinus Bonaparte, 1854, and
Sporadicus Cabanis & Heine, 1860; Riccordia has priority, and the type species in all is ricordii Gervais, 1835).
Ridgway (1911) and Cory (1918) recognized Riccordia for these species, but Peters (1945) merged it into
Chlorostilbon without comment. Also included in Riccordia is the recently extinct species bracei Lawrence, which
was probably a close relative of ricordii (Graves & Olson 1987). We include the extinct species R. elegans (Gould)
based upon Weller (1999) following R. bracei, but apparently doubts remain regarding the origin of the one extant
specimen as well as to its status as a distinct species or as a subspecies of bracei (Avibase 2012). Cyanophaia
bicolor (Gmelin, 1788) differs appreciably in plumage from the Riccordia species and the branch connecting it to
Riccordia is relatively long albeit not well-supported, such that one alternative might be to maintain it as a
monotypic genus, which could be supported by its apparent lack of well-developed tomial serrations (JVR & FGS,
pers. obs.). However, a precedent for not basing generic limits on tomial serrations occurs in the genus Polytmus in
the Polytminae, in which guainumbi (Pallas) has well-developed serrations (as do the other members of the
Polytminae) but theresiae (Da Silva Maia) does not (Stiles & Rico, submitted); nevertheless, the latter clearly must
be placed in Polytmus in the phylogeny (see Remsen et al. 2015). Schuchmann’s (1999) linear sequence placed
Cyanophaia following Panterpe and other Middle American species, although he suggested that it might be closer
to Chlorostilbon. The genetic data place C. bicolor in a strictly Antillean radiation with Riccordia; therefore, it is
not a relatively recent arrival from South or Middle America (cf. Bond 1959). We prefer to recognize the
relationship of bicolor and Riccordia by considering them congeneric. We note that Riccordia and Cyanophaia
were described on the same page by Reichenbach (1854); as first revisers, we choose Riccordia as the name of the
broader genus. Divergence in plumage color in bicolor, typical for many taxa on small islands, has masked its
relationship to Riccordia sensu stricto, which it replaces in the Lesser Antilles.
We also note that a comparable case exists in the Polytminae (Remsen et al. 2015). Two monospecific Lesser
Antillean genera, Eulampis and Sericotes, were found in the phylogeny to be embedded within Anthracothorax and
closely related to A. viridigula, which occurs widely in northeastern South America including Trinidad. Although
Bond (1959) considered Eulampis and Sericotes to be old Antillean endemics, a more recent colonization of the
Lesser Antilles by viridigula stock from Trinidad seems a more likely origin for them, followed by rapid
divergence in plumage and morphology on these small islands that has masked their relationships.
Subgroup A3 contains the other species of Cynanthus including its type, latirostris Swainson, 1827a, as well as
Chlorostilbon canivetii (Lesson) and salvini Cabanis & Heine. This relationship seems surprising at first sight
because in most aspects of their plumage, including a distinctive female plumage with a contrasting white
postocular stripe and deeply forked tails in the males, these two species resemble typical members of Chlorostilbon
and are less like the species of Cynanthus. The generic name Chloanges Heine, 1863, is available for these species,
which were distinguished from Chlorostilbon by the gray tips to the males’ rectrices. The type of Chloanges is
auriceps (Gould), of which canivetii and salvini were considered close relatives or subspecies until Howell (1993)
treated them as separate species (along with osberti Gould, 1860, which Howell also split from salvini). We have
verified identifications of the vouchers (Burke Museum) used by McGuire et al. (2014). However, given the
topology of the tree, recognizing Chloanges would render Cynanthus paraphyletic, as was also found by García-
Deras et al. (2008). We therefore propose that canivetii and salvini (and their close relatives) be included in

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Cynanthus Swainson, 1827a. This indicates that Cynanthus represents a northern Middle American radiation
distinct from the mainly South American subgroup A4, and that the “typical” plumage of Chlorostilbon is subject
to homoplasy. The southernmost member of the Chloanges group, canivetii, extends southward to northern Costa
Rica as a member of the Pacific coast dry forest avifauna of Middle America, whereas assimilis Lawrence, the
northernmost member of the southern radiation, is a member of the southwestern Costa Rica-Panama wet forest
avifauna, and hence is more similar ecologically to most South American species. Subgroup A4 comprises the bulk
of Chlorostilbon Gould,1853, including its type, mellisugus (Linnaeus, 1758), as determined by Zimmer (1950a).
The “green-tailed” members of Chlorostilbon, represented in the tree by C. poortmani (Bourcier), were formerly
separated in the genus Panychlora Cabanis & Heine, 1860, but the genetic data indicate that this species is firmly
nested within Chlorostilbon. Males of the other species have tails that are wholly dark steely-blue and vary from
nearly truncate (assimilis, mellisugus) to fairly deeply forked (e. g., gibsoni Fraser), but are never so deeply forked
as in the “Chloanges” species, now included in Cynanthus (see above).
Group B is also taxonomically polyphyletic (Fig. 1). Sister to the rest of the group (subgroup B1) are
“Hylocharis” leucotis (Vieillot) and “H.” xantusii (Lawrence). However, the phylogeny (McGuire et al. 2014)
separates the rest of Hylocharis Boie, 1831 including its type species sapphirina (Gmelin,1788) in group D (see
below). Ridgway (1911) and Cory (1918) treated leucotis and xantusii in the genus Basilinna Boie, 1831, but
Peters (1945) merged it into Hylocharis without comment. More recently, Howell & Webb (1995), Schuchmann
(1999) and Hernández et al. (2014) have provided evidence that Basilinna should be restored for leucotis (Vieillot,
1818), its type species, and xantusii, which is supported by the genetic data. Here again, the character used by
many authors to diagnose Hylocharis, the expanded red base of the bill in adult males, shows considerable
homoplasy and is of little phylogenetic value.
Subgroup B2 consists of two species of Campylopterus Swainson, 1827b (excellens Wetmore and rufus
(Lesson), on a long branch. The remainder of Campylopterus constitutes subgroup B4, which includes its type
species, largipennis (Boddaert, 1783). Hence, another name is required for excellens and rufus, both of which
inhabit northern Middle America. Of the available names, Pampa Reichenbach, 1854, has priority over
Sphenoproctus Cabanis & Heine, 1860; for both, the type species is curvipennis (Deppe, 1830), which is a close
relative of, and often considered conspecific with, excellens (e.g., Friedmann et al. 1950, Schuchmann 1999).
Pampa was recognized as a distinct genus in the 19
th
and early 20
th
centuries [e. g., by Ridgway (1911) and Cory
(1918)], but Peters (1945) subsumed it into Campylopterus, stating that the differences between Pampa and
Campylopterus were of “specific rather than generic value”. Although plumages of the species of Pampa do
resemble rather closely those of the sexually monomorphic species of Campylopterus, morphological differences
between the genera exist, as diagnosed by Ridgway (1911). Both Pampa species possess relatively longer and more
strongly graduated to cuneate tails vs. truncate to rounded tails in Campylopterus; the anterior toes are basally
syndactylous in Campylopterus but not in Pampa; the females in both genera have basically gray underparts, but
these are immaculate in Pampa but with green speckling laterally and sometimes with contrasting gorgets in
Campylopterus; the downy femoral tufts are more conspicuous in Campylopterus. A further difference may occur
in song: those of Pampa species appear to be much more elaborate and varied, whereas songs of those species in
Campylopterus with song descriptions are simpler and more repetitive (Hilty & Brown 1986; Stiles & Skutch
1989; Howell & Webb 1995; Hilty 2002; Schulenberg et al. 2007; González et al. 2011). Clearly, similarities in
plumage have masked considerable genetic and behavioral differences between these genera.
Subgroup B3 includes five genera with only one or two species each, all fairly distinctive and separated by
long branches: Klais Reichenbach, 1854; Abeillia Bonaparte, 1850a; Orthorhynchus Lacépède, 1799; Stephanoxis
Simon, 1897; and Anthocephala Cabanis & Heine, 1860. Given the heterogeneity of this group, we see no sensible
way to combine any two or three without having to include a quite dissimilar genus: for instance, combining the
two crested genera, Orthorhynchus and Stephanoxis, would also require including in the genus the very different-
looking Anthocephala. In contrast to most groupings in the Trochilini, biogeography produces no coherent pattern:
Orthorhynchus is Antillean, Abeillia is endemic to northern Middle America and Anthocephala to montane
Colombia, Klais ranges in the lowlands and foothills from Middle America south to central Bolivia and
Stephanoxis occurs in southeastern South America. Hence, the options are to combine all into an exceptionally
heterogeneous (by trochiline standards) genus Orthorhynchus Lacépède, 1799, or to leave them as a group of small
genera. The relatively long branches that unite them and the lack of morphological or biogeographic continuity
among them favor continued recognition of each genus, which also maintains taxonomic stability.

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FIGURE 1. The currently accepted classification of the Trochilini, with groups A, B, C and D and their subgroups, as derived
from the phylogeny; at the extreme right of each group are the new generic circumscriptions recommended and explained in the
text. See Appendix 2 for the complete generic reclassification of the Trochilini.
Cynanthus sordidus
Chlorostilbon ricordii
Chlorostilbon swainsonii
Chlorostilbon maugaeus
Cyanophaia bicolor
Cynanthus latirostris
Cynanthus latirostris doubledayi
Chlorostilbon canivetii salvini
Chlorostilbon canivetii
Chlorostilbon mellisugus
Chlorostilbon lucidus lucidus
Chlorostilbon lucidus pucherani
Chlorostilbon poortmani
Chlorostilbon assimilis
Chlorostilbon melanorhynchus
Phaeoptila
Riccordia
Cynanthus
Chlorostilbon
Hylocharis xantusii
Hylocharis leucotis
Campylopterus rufus
Campylopterus excellens
Klais guimeti
Abeillia abeillei
Orthorhyncus cristatus
Stephanoxis lalandi
Campylopterus villaviscensio
Campylopterus ensipennis
Campylopterus falcatus
Campylopterus largipennis
Campylopterus duidae
Campylopterus hyperythrus
Campylopterus hemileucurus
Group A
Basilinna
Pampa
Klais
Abeillia
Orthorhynchus
Stephanoxis
Anthocephala
Campylopterus
Group B
Microchera albocoronata
Elvira chionura
Elvira cupreiceps
Goethalsia bella
Goldmania violiceps
Thalurania ridgwayi
Eupherusa poliocerca
Eupherusa cyanophrys
Eupherusa eximia
Eupherusa nigriventris
Chalybura urochrysia
Chalybura buffonii
Thalurania glaucopis
Thalurania watertonii
Thalurania furcata
Thalurania furcata
Thalurania colombica colombica
Thalurania colombica fannyae
Thalurania colombica fannyae
Thalurania colombica colombica
Group C
Microchera
Goldmania
Eupherusa
Thalurania
Chalybura
Phaeochroa cuvierii
Leucippus fallax
Leucippus taczanowskii
Leucippus baeri
Taphrospilus hypostictus
Eupetomena macroura
Aphantochroa cirrochloris
Leucippus chlorocercus
Trochilus polytmus
Trochilus polytmus
Trochilus scitulus
Trochilus scitulus
Trochilus polytmus
Trochilus scitulus
Amazilia viridifrons wagneri
Amazilia violiceps
Amazilia viridifrons
Amazilia cyanocephala
Amazilia saucerrottei hoffmanni
Amazilia beryllina
Amazilia cyanura
Amazilia edward
Amazilia saucerrottei
Amazilia castaneiventris
Amazilia tobaci
Amazilia tobaci
Amazilia viridigaster
Amazilia rutila
Amazilia yucatanensis
Amazilia tzacatl handleyi
Amazilia tzacatl
Amazilia amazilia
Amazilia franciae
Lepidopyga goudoti
Chrysuronia oenone
Amazilia versicolor
Lepidopyga coeruleogularis
Hylocharis grayi
Amazilia brevir. chionopectus
Amazilia brevirostris
Amazilia leucogaster
Leucochloris albicollis
Amazilia lactea
Hylocharis sapphirina
Hylocharis chrysura
Amazilia chionogaster
Amazilia viridicauda
Amazilia rosenbergi
Amazilia decora
Amazilia amabilis
Amazilia candida
Hylocharis eliciae
Hylocharis cyanus
Juliamyia julie
Chlorestes notatus
Group D
Phaeochroa
Leucippus
Thaumasius
Taphrospilus
Eupetomena
Talaphorus
Trochilus
Leucolia
Saucerottia
Amazilia
Amazilis
Uranomitra
Chrysuronia
Leucochloris
Chionomesa
Hylocharis
Elliotia
Polyerata
Chlorestes
Trochilini (Emeralds)
1
2
3
4
1
2
3
4
1
2
1
7
2
3
4
5
6

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Subgroup B4 includes the remaining species in Campylopterus Swainson, 1827b. Although some of the branch
lengths are rather long, we see nothing to be gained by splitting Campylopterus into three or four small genera (at
least one of which could require a new name), and recommend leaving all of the species in Campylopterus, which
also maintains taxonomic stability. We note that the potential problem with the C. rufus sample in this group
(McGuire et al.2014) was due to a mislabeled sample of C. hemileucurus (Deppe) (J. Bates, personal
communication).
Group C (Fig. 1) is characterized by type A tomial serrations; the topology of the genetic tree indicates that
such serrations were derived independently from those of clade A (see above). This group breaks cleanly into two
subgroups (Fig. 1). Subgroup C1 includes four more or less distinct groupings: (1) Microchera Gould, 1858 and
Elvira Mulsant et al., 1866; (2) Goldmania Nelson, 1911 and Goethalsia Nelson, 1912; (3) Thalurania ridgwayi
Nelson 1900; and (4) Eupherusa Gould, 1857. Given the very close relationship indicated by the branch lengths
that join them, we favor combining Microchera and Elvira into a single genus: Microchera has priority. Monotypic
Microchera albocoronata (Lawrence, 1855) has been recognized traditionally by the shining white crown and
purplish underparts of the males. Although Salvin (1892), Cory (1918) and Simon (1921) recognized its close
relationship with Elvira, Schuchmann (1999) placed 16 genera between Microchera and Elvira in his linear
sequence. All three species are small hummingbirds with extensive white in the tail, and all occur in foothill and
montane regions of southern Middle America. Moreover, the female of Microchera is much less distinctive, and its
bronzy uppertail coverts and white in the tail resemble those of female Elvira.
For the second group, we also see no reason not to combine Goethalsia and Goldmania: they share an unusual
type of undertail coverts (fluffy with stiff rachises) and a restricted parapatric distribution in small, isolated
highlands in eastern Panama and adjacent Colombia. They differ mainly in the distribution of contrasting plumage
areas on the otherwise mostly green body. Their close relationship has always been suspected, as signaled by their
adjacent placement in all published linear sequences. Goldmania Nelson, 1911 has priority over Goethalsia
Nelson, 1912.
The surprise in this group is Thalurania ridgwayi, which is clearly separated from the rest of Thalurania
Gould, 1848 (which is in subgroup C2, separated by a long branch). As far as we can tell, ridgwayi has always been
included in Thalurania since its description by Nelson in 1900, and was considered to form a superspecies with T.
colombica by Schuchmann (1999). Its violet-blue crown, iridescent green throat and steely blue tail do resemble
the plumages of other male Thalurania, although the crown is more bluish and much more extensive; however,
ridgwayi differs from other species of Thalurania in its greenish-black underparts and much less deeply forked tail.
Closest to ridgwayi and separated by a short branch is the genus Eupherusa Gould, 1857, with four closely related
species. Males of ridgwayi differ from those of all species of Eupherusa in lacking white in the tail and rufous in
the remiges. However, the blue crown of male ridgwayi is quite similar to that of E. cyanophrys Rowley & Orr, and
the blackish underparts are shared with E. nigriventris Lawrence, which also has less extensive rufous in the
remiges; its shallowly forked tail also resembles more closely those of Eupherusa species. Females of ridgwayi and
Eupherusa share gray underparts and differ mainly in tail patterns, although here again E. nigriventris has less
extensive white in the tail than do those of the other Eupherusa species. Biogeographically, ridgwayi has a Pacific
slope distribution, as do three species of Eupherusa. Including ridgwayi in Eupherusa would constrict the northern
limit of Thalurania to the humid Caribbean lowlands of Guatemala to northeastern Honduras. Although fieldwork
on ridgwayi is needed to reveal whether it shares vocal and behavioral similarities to Eupherusa and differs from
true Thalurania, we consider that the available evidence favors the inclusion of ridgwayi in the genus Eupherusa.
Subgroup C2 includes only two genera, Chalybura Reichenbach, 1854, and Thalurania Gould, 1848, including
its type, furcata (Gmelin, 1788). Thalurania and Chalybura have never been considered closely related, much less
sister genera; for example, 13 genera separate the two in Schuchmann’s sequence. However, they share some
plumage features heretofore unremarked: (1) the long white undertail coverts of most Chalybura are shared with
Thalurania furcata baeri Hellmayr; (2) the somewhat blotchy iridescent green throats of most Chalybura recall
that of T. glaucopis (Gmelin); and (3) the steely blue tail of C. buffonii (Lesson) is similar to that of Thalurania.
Males of both genera are notably aggressive and territorial at rich aggregations of flowers. The two genera differ
vocally: male Chalybura sing and are highly vocal in aggressive interactions, whereas males of Thalurania are
relatively silent (Stiles & Skutch 1989; FGS, pers. obs.). Moreover, Thalurania species are much smaller, and their
males have much more deeply forked tails than do those of Chalybura. The two genera are separated by a long
branch; hence, we recommend continuing recognition of them as separate genera.

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By far the most diverse and nomenclaturally complicated group of the emeralds is group D (Fig. 1), which
includes all of the problems associated with genera such as Leucippus Bonaparte, 1850a and Amazilia Lesson,
1843 (dubbed the “amazilian complex” in Stiles et al. 2017), among others. This group can be broken reasonably
well into seven subgroups. Subgroup D1 contains only two species: Phaeochroa cuvierii (De Lattre & Bourcier,
1846) and Leucippus fallax (Bourcier, 1843b). Phaeochroa cuvierii differs greatly in size from L. fallax and also in
plumage pattern, with speckled-scaly underparts and conspicuously white-tipped rectrices; in contrast, L. fallax is
uniform buff below with weakly whitish-tipped rectrices. Phaeochroa cuvierii also has a louder, much more
elaborate song than that of L. fallax, and the adult males have the rachises of the outermost primaries thickened and
flattened, rather like those of Campylopterus. Given the long branch lengths to the two species, we consider that
Phaeochroa Gould, 1861, and Leucippus Bonaparte, 1850a (for which fallax is the type, as fixed by Gray 1855) are
best left as monotypic genera. The genetic data also clearly reject the inclusion of Phaeochroa in Campylopterus,
as was done by Schuchmann (1999); the “saber” wings of each have evolved independently. Phaeochroa cuvierii
occurs on the Caribbean slope of Middle America from southern Mexico and Belize to northern Colombia, as well
as on the Pacific slope from northwestern Costa Rica to eastern Panama; L. fallax occupies the arid Caribbean
coastal lowlands of northern Colombia and Venezuela. The two are nearly parapatric in Colombia, with P. cuvierii
occurring in moister sites further inland. We have noted (Stiles et al. 2017) that Bonaparte (1854) separated fallax
in the monotypic genus Doleromya Bonaparte, 1854 before Gray’s fixation of it as the type species of Leucippus,
but because the older genus takes precedence, Doleromya is an objective synonym of Leucippus.
Subgroup D2 includes two groups: “Leucippus” taczanowskii Sclater and “L.” baeri Simon, and the
monotypic genera Aphantochroa, Eupetomena and Taphrospilus. The first group has usually been included in
Leucippus in the past due to the whitish underparts of the two species, but this is clearly untenable because the type
of that genus is fallax. Chubb’s (1916) rejection of the generic name Thaumasius Sclater, 1879 was unjustified
(Stiles et al. 2017), so this is the appropriate generic name, and taczanowskii Sclater, 1879 is the type species by
monotypy. Both Thaumasius species are sexually monomorphic and are unusually dull, brownish species quite
similar in plumage; they have small, allopatric ranges centered in arid areas in western Peru: baeri in the
Tumbesian region, taczanowskii in the upper Marañón valley. Taken together, their distribution pattern is shared by
many bird taxa in the region; these two fit the classic requirements of a superspecies.
Rather long branches connect the three species of the second group, currently placed in three monotypic
genera: Eupetomena Gould, 1852; Aphantochroa Gould, 1853; and Taphro s p i lu s Simon,1910; a situation
reminiscent of subclade B3 above. All three are rather large hummingbirds. Aphantochroa cirrochloris (Vieillot,
1818) and Eupetomena macroura (Gmelin, 1788) are sympatric in southeastern Brazil, whereas Taphrospilus
hypostictus (Gould, 1862) occurs on the lower eastern slope of the Andes from southern Colombia to northern
Argentina. All three show little or no sexual dimorphism in color, as in Thaumasius; the plumages of Thaumasius
and hypostictus are sufficiently similar that Schuchmann (1999) had placed all three species in Leucippus.
Although hypostictus was originally described in Aphantochroa, it is somewhat brighter in coloration than A.
cirrochloris (fairly bright green-and-white vs. dull green-and-gray underparts). Both sexes of Eupetomena are dark
blue with deeply forked tails (more deeply in males). Males of Aphantochroa and Eupetomena share broad,
flattened rachises of the outer primaries, suggestive of the “saber” wings of Campylopterus of clade B. Indeed,
Schuchmann (1999) had placed both species in Campylopterus because of this, but genetic data strongly refute that
relationship. Taphrospilus differs ecologically and geographically from these two, and its males lack “saber-type”
wings. For these genera, information regarding branching pattern, branch lengths, biogeography and phenotypic
differentiation is discordant, leaving three options for our generic rearrangement: (1) include all three in a single
heterogeneous genus, for which Eupetomena has priority; (2) lump Aphantochroa into Eupetomena (which are
sisters in the phylogeny); or (3) continue to recognize the three monotypic genera. We consider option 1 the worst
alternative because the long branch lengths mask considerable genetic and phenotypic diversity while ignoring
intrageneric relationships. At the other extreme, option 3 also ignores these relationships as well as patterns of
phenotypic variation; its principal advantage lies in maintaining nomenclatural stability. Option 2 would represent
a compromise between these different criteria: it would recognize the sister status of macroura and cirrochloris as
well as their shared biogeography and their sexually dimorphic wings, but would override the shared dull
coloration of hyposticta and cirrochloris and the sharp contrast in coloration and tail shape between the latter and
macroura. However, the phylogeny provides multiple precedents for doing so. Similar dull plumages without
bright colors or sexual dimorphism due to conservative plumage evolution or homoplasy occur in several groups of

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the amazilian complex and the broadly circumscribed Leucippus in the traditional classification, while the genetic
tree indicates that some new generic groupings (q.v., Cynanthus, Chrysuronia and Chlorestes) include species
differing widely in colors and patterns of plumage, sexual dichromatism and tail shapes. Moreover, many of the
new generic groupings clearly show geographic congruence. We therefore conclude that option 2 is the most
informative regarding relationships and recommend lumping Aphantochroa into Eupetomena while maintaining a
monotypic Taphro s p i l u s .
Subgroup D3 includes three groups. The most distinct, a clear outlier perhaps constituting a subgroup by itself,
is “Leucippus” chlorocercus Gould, 1866, which clearly belongs in the monotypic genus Talaphorus Mulsant & E.
Verreaux, 1874, of which it is the type species). This species is fairly similar in plumage to the two species of
Thaumasius, neither of which had been named when Talaphorus was described. It differs most obviously from
Thaumasius in having gray lateral rectrices with a pronounced subterminal blackish band and in its more grayish-
white rather than brownish-white underparts, and also in ecology and biogeography, being restricted to the banks
and islands of the larger rivers in the humid upper Amazon basin. It provides another example of a species whose
genetic distinctiveness was masked by conservative plumage evolution and the absence of strong patterns and
bright colors. The second subgroup includes only the genus Trochil us, a Jamaican endemic; the morphological and
geographical distinctiveness and position in the tree of this genus clearly justify its continued recognition. The only
question raised by the phylogeny with respect to Trochi lus is whether the recognition of two species as in
Schuchmann (1999) is justified, because the genetic samples of polytmus (Linnaeus, 1758) and scitulus (Brewster
& Bangs) are intermixed for the loci sampled. However, Graves (2015), who presented a detailed genetic and
biogeographic analysis that established that polytmus and scitulus are nearly parapatric with a very narrow zone of
limited hybridization and introgression, recommended species rank for both forms, a recommendation followed
here.
The third group of subgroup D3 includes two well-defined sections of the amazilian complex. The first
includes the taxa violiceps (Gould), viridifrons (Elliot), and wagneri Phillips, the latter often being considered a
subspecies of viridifrons although the genetic data suggest that it is less close to viridifrons than the latter is to
violiceps. For these, finding an available generic name has proven more difficult. In the past, these species have
been included in Cyanomyia Bonaparte, 1854, Uranomitra Reichenbach, 1854, Agyrtria Reichenbach, 1854,
Agyrtrina Chubb, 1916, Amizilis Gray, 1840, or Amazilia Lesson, 1843. Of these, Agyrtria may be eliminated
because it is a synonym of Polytmus Brisson, 1760, and Amizilis is a synonym of Cynanthus (see Stiles et al. 2017).
The type species of all of the other genera fall out in different parts of the phylogeny. The type species of
Cyanomyia, which is cyanocephala (Lesson, 1829), is now included in Saucerottia Bonaparte, 1850b (see next
section), and thus Cyanomyia is a synonym of Saucerottia. The type species of Uranomitra, franciae (Bourcier &
Mulsant, 1846), is in subgroup D5 as is amazilia (Lesson & Garnot, 1827), the type of Amazilis (see Stiles et al.
2017 and below); Amazilia sensu stricto constitutes the subgroup D4; most species of Agyrtrina are in subgroup
D5. This leaves Leucolia Mulsant et al., 1866, within which they considered Cyanomyia and Leucolia to be
subgenera; they included violiceps in the former subgenus. The genus Leucolia sensu stricto included viridifrons as
well as four other species that in the tree are included in other genera. This leaves viridifrons eligible as the type
and indeed, it was tentatively so designated by Elliot (1879). Therefore, we recommend recognizing the genus
Leucolia, and we fix viridifrons (Elliot, 1871) as its type species; this also includes transferring violiceps and
wagneri from Cyanomyia to Leucolia. Rodríguez-Gómez & Ornelas (2015) further documented the close genetic
relationship between violiceps and viridifrons.
The second section includes the species of Saucerottia Bonaparte, 1850b with its type species saucerottei (De
Lattre & Bourcier, 1846), which the genetic data clearly support ranking as a genus. Many authors recognized
Saucerottia as a genus until Peters (1945) demoted it to a subgenus of Amazilia. Subsequent classifications
followed Peters until Schuchmann (1999) resurrected it as a genus. McGuire et al. (2014) found a similar group
composition except that the species castaneiventris and cyanocephala were also members of Saucerottia.
Schuchmann (1999) had placed the former in Amazilia because of its rufous belly, and the latter in Agyrtria due to
its mostly white underparts – yet another warning that coloration is not a dependable guide to relationships in the
Trochilini! All species of Saucerottia measured by FGS (unpubl. data) differ from the rest of the amazilian
complex in their narrower, more pointed wings, and all share notably bright, glittering green feathers on the throat
and breast or over the entire underparts. All species of Saucerottia occur in Middle America and northern South
America. Also of interest is that the genetic data justify the separation of hoffmanni (Cabanis & Heine) of southern

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Mesoamerica from the South American saucerottei at the species level because they are not sister taxa in the tree;
most older classifications had also made this separation (although hoffmanni was often treated under the species
epithet sophiae) until Peters (1945) lumped it into saucerottei. FGS (in Stiles & Skutch 1989) had long suspected
that hoffmanni deserved species rank due to the range disjunction (northern Costa Rica to northern Colombia),
pronounced differences in song, and lesser differences in plumage (but on a par with differences between other
species in this genus, e.g., S. saucerottei and S. tobaci). Jiménez & Ornelas (2016) confirmed the genetic
relationship of hoffmanni with a complex of Middle American species distinct from the South American
saucerottei group. However, they also used the species name sophiae, which Hellmayr (1913) had decisively
shown to be a synonym of S. saucerottei warsewiczii; Hellmayr therefore used hoffmanni for the Middle American
species. This was followed by Peters (1945) and Schuchmann (1999).
Subgroup D4 includes three species: rutila (De Lattre), yucatanensis (Cabot) and tzacatl (De la Llave). These
three should constitute the restricted genus Amazilia Lesson, 1843 with rutila (De Lattre, 1843) as its type species;
see Stiles et al. 2017). The genetic distinctiveness of this group had long been masked by similarities in plumage
with some other members of the amazilian complex. However, all three species have conspicuous rufous tails;
buffy bellies are shared by rutila and yucatanensis, and bright green breasts by yucatanensis and tzacatl. They are
among the northernmost members of the amazilian complex: yucatanensis breeds as far north as central Texas,
rutila occurs mainly on the Pacific slope from Mexico to Costa Rica, and tzacatl on the humid Atlantic slope of
Mesoamerica from Belize southward, on both slopes in Panama and northern Colombia and south on the humid
Pacific slope to northern Ecuador.
Subgroup D5 is more complicated than D4. This subgroup contains two successive outliers on long branches,
then a compact group of seven species weakly divided into two groups. The first outlier is “Amazilia” amazilia
(Lesson & Garnot, 1827), which we argued (Stiles et al. 2017) should be placed in the genus Amazilis Gray, 1855,
of which it is the type species. This species appears to be an old isolate on the Pacific slope of Ecuador and Peru, to
judge from the distinctiveness in plumage of its several subspecies; one of these, alticola Gould, was accorded
specific rank by Schuchmann (1999) because of possible sympatry without hybridization with other subspecies in
extreme southwestern Ecuador. However, Krabbe & Ridgely (2010) analyzed the biogeography of this group in
more detail and recommended that alticola be retained as a subspecies of amazilia, and this is followed here. The
plumage pattern of amazilia, containing brilliant green, white and rufous areas is also unique in the amazilian
complex.
The second outlier is “Amazilia” franciae (Bourcier & Mulsant, 1846), which we recommend be treated in the
monotypic genus Uranomitra Reichenbach, 1854, of which it is the type, named by Elliot (1879). However, type
fixation of Uranomitra also presents complications. Gray (1855) had fixed the type as Trochilus quadricolor
(Vieillot, 1823) but, as stated by Salvin (1892), Vieillot (1823) named the “colibri quadricolor” as Trochilus
quadricolor on p. 555, and his description clearly applied to a species of “Lampornis” (=Anthracothorax in current
classifications), and on p. 573 in the same work he also named the “oiseau-mouche quadricolor” with the name
Trochilus quadricolor for an emerald, perhaps franciae, but this quadricolor is clearly a junior homonym and thus
is not available. Ridgway (1911) gave the type of Uranomitra as “Tr. quadricolor, i.e., Tr. verticalis Deppe, 1830”.
However, Hellmayr (1913) showed that verticalis was a synonym of T. cyanocephala Lesson (now in Saucerottia
Bonaparte, 1850b), and thus is also not applicable. Therefore, the only applicable type designation for Uranomitra
is that by Elliot (1879) of franciae, which we accept here. This species had been placed in Agyrtria by many
authors, most recently including Schuchmann (1999). However, its plumage is brighter than those of other white-
bellied emeralds, and it also shows some sexual dichromatism with females having duller blue crowns than do
males. This species also differs from others in the amazilian complex in its basically Andean distribution, and
morphologically from those species measured by FGS (unpubl. data) by its longer, slightly thicker bill, longer tarsi
and larger feet. It is also one of the heaviest species in this complex and has the highest wing loading. At the
southern extreme of its distribution franciae, (as the subspecies cyanocollis Gould, possibly recognizable as a
distinct species) is limited to middle elevations of the dry Marañón valley in Peru, where it may approach parapatry
with Amazilis amazilia, which occupies similar habitats on the Pacific slope (Schulenberg et al. 2007).
The rest of subgroup D5 breaks into two smaller, compact subgroups, separated by a very short branch. The
first consists of three species in three currently recognized genera: Lepidopyga goudoti (Bourcier, 1843), the type
of this genus; Chrysuronia oenone (Lesson, 1832a), also its type; and “Amazilia” versicolor (Vieillot,1818). The
second group includes four species in three genera: Lepidopyga coeruleogularis (Gould); Hylocharis grayi (De

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Lattre & Bourcier); Agyrtrina brevirostris (Lesson); and “A.” leucogaster (Gmelin). Nowhere in the Trochilini is
the mismatch between plumage features and phylogeny so striking. Two main options exist here: 1) combine all
seven species in a single genus, for which Chrysuronia Bonaparte, 1850a takes priority over Lepidopyga
Reichenbach,1854, Agyrtrina Chubb, 1916, and Eucephala Reichenbach, 1854 (currently subsumed in Hylocharis
Boie, whose type species, sapphirina, is in subgroup D6); or 2) separate the two groups in two genera (respecting
branching pattern but ignoring branch lengths). Chrysuronia has priority for the first group, Eucephala with grayi
(De Lattre & Bourcier, 1846) as its type for the second. We favor the first option, given the very short branch
separating the two groups. Adding the latter four species to Chrysuronia would produce a genus scarcely more
heterogeneous in colors and patterns than that with only the first three. Moreover, this would unite in a single genus
the species in two genera, Lepidopyga and Agyrtrina, which must disappear in the interests of priority. A third
option would be to include all of subgroup D5 in a single heterogeneous genus Amazilis Gray, 1855, which we
consider the least acceptable because it ignores the decidedly longer branch lengths to amazilia and franciae.
Although heterogeneous, a broad Chrysuronia does include some common themes: the females of all are basically
white below with green speckling laterally, as are the males of those species not showing sexual dichromatism.
Males of most of the dimorphic species show conspicuous blue on the head or breast; the most divergent in form
are the two former species of Lepidopyga with their more deeply forked tails with narrow outer rectrices, which
clearly represents homoplasy because they are not sisters in the tree, nor even members of the same group within
Chrysuronia.
Subgroup D6 breaks up into four groups separated by moderate branch lengths. The first group, sister to the
rest in the tree, includes only albicollis (Vieillot,1818), type of the genus Leucochloris Reichenbach, 1854, and we
favor continued recognition of this monotypic genus. Its plumage pattern is unique: a clear white throat, green
breast, white belly and conspicuous white in the tail. The second group consists of the two species “Amazilia”
fimbriata (Gmelin) and “A.” lactea (Lesson), which share a similar plumage pattern, with a medial white streak
extending anteriorly from the belly; they differ mainly in having green vs. blue anteriorly and on the sides. These
species have been placed in Agyrtria or Polyerata by most recent authors (when not in Amazilia itself). Elliot
(1879) had named lactea as the type species of Cyanochloris Reichenbach, 1854 but he overlooked the earlier
designation by Gray (1855) of Trochilus coeruleogularis Gould, 1847 as the type species of Cyanochloris. Because
coeruleogularis is now considered a subspecies of Chalybura buffonii (Lesson), Cyanochloris is a synonym of
Chalybura Reichenbach, 1854. However, the name Chionomesa Simon, 1921 is available; in fact, Simon included
in it only these two species (although considering as species several subspecies of fimbriata in particular), but did
not name a type species. We therefore recommend recognizing Chionomesa for lactea and fimbriata, and fix lactea
(Lesson, 1833) as its type species. We note here that Lesson’s name lactea was first proposed in the Index
Générale, the final part of his third major work, Les Trochilidées (1832b-1833) and dates from 1833, not 1829 (in
which the description and plate were published in Lesson’s earlier work (1829-1830) under the name “Le Saphir,
Ornismya sapphirina”. Lesson (1843) proposed the name lactea for that part of the description and the plate,
recognizing that the rest applied to sapphirina Gmelin (see below).
The third group of D6 includes Hylocharis sapphirina (Gmelin, 1788) (type of the genus Hylocharis Boie,
1831) and H. chrysura (Shaw); these two therefore constitute the restricted genus Hylocharis. They share the
expanded red base of the bill formerly used to distinguish this genus, but this character is clearly subject to
homoplasy because it appears in several species placed elsewhere by the genetic data. Its two species are quite
different in color and pattern but share one unique feature: the contrasting rufous chin and upper throat. Both occur
in southern Brazil and adjacent countries, but sapphirina also has a broad, disjunct range in eastern Colombia,
southern Venezuela, the Guianas, and northern Brazil.
The fourth group consists of the species “Amazilia” chionogaster (von Tschudi), and “A.” viridicauda
(Berlepsch), which usually had been included in Leucippus in the past (see Stiles et al. 2017); in fact, chionogaster
was named the type species of Leucippus by Elliot (1879) when fallax was segregated by some authors in the
monotypic genus Doleromya Bonaparte, 1854 (see above). Their inclusion in Amazilia by Zimmer (1950b) was
based on the similarity in plumage between them and A. candida and A. versicolor. Meyer de Schauensee (1966)
and most subsequent authors except Schuchmann (1999) have followed Zimmer’s treatment. However, the tree
demonstrates that this type of coloration is clearly subject to much homoplasy and that these species cannot be
included in either Amazilia or Leucippus. Neither we nor Peters (1945:59) have found any other generic name
proposed specifically for them. We also cannot find support for the statement by Schuchmann (1999) that these

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species had been “often alternatively placed in genus Chionogaster”, or indeed that the name Chionogaster has
ever been used in a generic sense: it does not appear in Zoonomen (Peterson 2009) or in any of the synonymies of
the original sources we have consulted. A new generic name for chionogaster and viridicauda is therefore
necessary. We propose
Elliotia, gen. nov.
Included species: Trochilus chionogaster von Tschudi, and Leucippus viridicauda Berlepsch. Both species weigh
4.5–5.5g, lack sexual dichromatism, and have mostly white underparts speckled with green laterally; they differ in
the amount of white in the tail. Both occur on the lower eastern slope of the Andes in central Peru (viridicauda) and
from northern Peru to northwestern Argentina (chionogaster). We select chionogaster (von Tschudi, 1846) as the
type species. The genus name honors Daniel Giraud Elliot for his important early contributions to clarifying the
generic taxonomy of the Trochilidae (see Stiles et al. 2017).
Subgroup D7 breaks up into two clear-cut groups. The first consists of the species amabilis (Gould), decora
(Salvin) and rosenbergi Boucard, which may be allocated to the genus Polyerata Heine, 1863; its type species is
amabilis (Gould, 1851). Weller (2000) also included several other species in this genus, but all are placed in other
sections of the phylogeny. The second group includes five species in four genera, arranged in a stepwise “cascade”
with very short branches separating them, such that dividing the group into two to four genera (at least one of
which would require a new generic name) would be arbitrary; thus, we treat all five species as congeneric. For this
genus, Chlorestes Reichenbach,1854 has priority over Juliamyia Bonaparte, 1854, its type species being Trochilus
notatus (Reich,1795). Damophila, published by Reichenbach in the same work and on the same page as Chlorestes,
is not available because it is preoccupied (Özdikmen 2008, see Stiles et al., 2017); Özdikmen’s suggested
replacement name Neodamophila is therefore a synonym of Chlorestes should julie (Bourcier, 1842), the type
species of Juliamyia, and notatus be considered congeners, as we recommend here. All five are small
hummingbirds (<4g in mass), with nearly truncate to strongly rounded tails in both sexes, differing mainly in colors
and patterns; all but candida share moderate to strong sexual dichromatism. We also note that the genetic data
strongly refute the inclusion of notatus in Chlorostilbon, as was done by Schuchmann (1999) because of the
similarity of the males’ plumage colors. In fact, Chlorostilbon is among the most distantly related genera in the
Trochilini (McGuire et al. 2014).
Finally, we leave unclassified two Middle American species for which no genetic samples were available to
McGuire et al. (2014): “Amazilia” boucardi (Mulsant,1877) and “A.” luciae (Lawrence, 1868). The former was
treated in the monotypic genus Arenella by Simon (1921), and as a member of Lepidopyga by Cory (1918) and
Ridgway (1911), while luciae was described in Thaumatias by Lawrence. Both species were placed in Amazilia by
Peters (1945) and most subsequent authors. Schuchmann (1999) and Weller (2000) included them in Polyerata and
considered them to comprise a superspecies, although they differ appreciably in plumage: boucardi bears some
resemblance to “Lepidopyga” goudoti, including in that the outer rectrices of the males are similarly narrow,
whereas the plumage of luciae resembles more that of Polyerata. However, given the frequently unreliable nature
of plumage color and pattern as generic characters among the Trochilinae, we prefer to treat both as incertae sedis
until genetic data become available.
As outlined above, allocating species to genera among the emeralds has been fraught with difficulties. While
we would like to preserve diagnosability of genera and retain what is salvageable of existing nomenclature in the
interests of stability, our goal has been to bring the results into as close a concordance as possible with the
branching pattern and branch lengths in the genetic tree (McGuire et al. 2014). Where no clear recommendation
was possible, we have clearly presented the options. To help clarify the latter, we present a list of the generic names
mentioned above, when and in what form they were proposed, their type species, and when and by whom these had
been fixed in Appendix 1, and we summarize our recommendations as a revised list of genera and species of the
Trochilini in Appendix 2.
The generic rearrangements we recommend here based upon the phylogeny of McGuire et al. (2014) represent
a drastic change from the three most recent classifications of the Trochilinae by Peters (1945), Schuchmann (1999)
and Dickinson & Remsen (2013) in numbers of species per genus (summarized in Table 1) and generic

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circumscriptions (Fig. 1, Appendix 2). The classification by Peters (1945) was characterized by widespread
lumping at both the species and genus levels, although he retained various species found to be synonyms or hybrids
by later authors. Most of the changes since are due to the breakup of several large genera (especially Chlorostilbon,
Amazilia and Hylocharis) into smaller genera or through reassigning some species into previously small or
monotypic genera (notably Chrysuronia and Chlorestes), as well as the elimination of several genera for reasons of
priority. The increase in the number of species by Schuchmann (1999) reflect rejection of several of Peters’
unsubstantiated lumpings; the increase by Dickinson & Remsen (2013) is mostly due to the restoration of species
status to several species lumped by Schuchmann into Chlorostilbon mellisugus but recognized elsewhere,
including those that the phylogeny mandates transferring to the genus Riccordia. The increase to 110 species in the
classification presented here is due to recent recognition of species status for distinctive subspecies in
Anthocephala and Stephanoxis and the inclusion of two recently extinct species of Riccordia.
TABLE 1. Changes in the numbers of genera, species and species per genus in the four most recent classifications of the
Trochilini.
1
= These genera were subdivided into subgenera.
Acknowledgments
McGuire’s research was supported by National Science Foundation grants DEB 0330750 and DEB 0543556. We
thank Robert C. Faucett for photographing vouchers at the Burke Museum, University of Washington and J. M.
Bates for checking vouchers the Field Museum of Natural History. Vitor Piacentini, Alan Peterson, Richard
Schodde, Dan Lane, Terry Chesser and Frank Steinheimer provided helpful discussions and comments on the
manuscript. Finally, we are grateful to Pamela Rasmussen for her painstaking editing of the manuscript.
Number of species per genus Peters (1945) Schuchmann (1999) Dickinson & Remsen (2013) Present study
117121510
245511
33115
41122
50112
60301
70100
81111
91
1
10 1
10 0 0 0 2
11 0 1 1 0
12 0 0 0 0
13 0 1 0 0
14 1 0 0 0
18 0 0 1 0
27 1
1
00 0
29 0 0 1 0
Incertae sedis 0 0 0 2
Total species 96 100 106 110
Total genera 29 28 28 35

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Literature cited
Avibase (2012) The world bird database: factsheet of Chlorostilbon elegans. Available from: https://avibase.bsc-eoc.org/
avibase.jsp?lang=EN (accessed 17 October 2017)
Boddaert, P. (1783) Table des Planches Enluminez d’Histoire Naturelle, de M. D’Aubenton avec les Dénominations de M.M.
Buffon, Brisson, Edwards, Linnaeus et Latham, précède d’un Notice des Principaux Ouvrages Luminées. Unknown
publisher, Utrecht, 67 pp.
Boie, F. (1831) Über Species und einige ornithologische Familien und Sippen. Isis von Oken, 24 (4), Cols. 542–548.
Bonaparte, C.L. (1850a) Conspectus Generum Avium. Vol. 1. E.J. Brill, Lugduni Batavorum, 543 pp.
Bonaparte, C.L. (1850b) Note sur les Trochilidés. Comptes Rendus hebdomadaires de l’Académie des Sciences de Paris, 30,
379–383.
Bonaparte, C.L. (1854) Tableau des oiseaux-mouches. Revue et Magasin de Zoologie pure et appliquée, 6, 248–257.
Bond, J. (1959) Derivations of Lesser Antillean birds. Proceedings of the Academy of Natural Sciences of Philadelphia, 131,
89–103.
Boucard, A. (1893) Description of supposed new species of humming birds in Boucard’s museum. The Humming Bird, 3 (1),
6–10.
Bourcier, J. (1842) Descriptions de trois nouvelles espèces de oiseaux-mouches. Revue Zoologique, 1842, 373.
Bourcier, J. (1843) Oiseaux-mouches nouveaux ou mal connus. Revue Zoologique, 1843, 70–72.
Bourcier, J. & Mulsant, E. (1943) Descriptions et figures de plusieurs espèces nouvelles d’oiseaux-mouches. Annales des
Sciences Physiques et Naturelles, d’Agriculture et d’Industrie de Lyon, 6, 36–49.
Bourcier, J. & Mulsant, E. (1846) Description de vingt espèces nouvelles d’oiseaux-mouches. Annales des Sciences Physiques
et Naturelles, d’Agriculture et d’Industrie de Lyon, 9, 312–332.
Cabanis, J. & Heine, F. (1860) Verzeichnis der Ornithologischen Sammlung, mit kritischer Anmerkungen und Beschreibung
der neuen Arten systematisch bearbeitet. Museum Heineanum, 3, 3–81.
Caverzare, V., Silveira, L.F., De Vasconcelos, M.F., Grantasu, R. & Straube, F.C. (2014) Taxonomy and biogeography of
Stephanoxis Simon 1897 (Aves: Trochilidae). Papeis Avulsos de Zoologia, 54, 69–79.
https://doi.org/10.1590/0031-1049.2014.54.07
Chubb, C. (1916) The Birds of British Guiana. Vol. 1. Bernard Quaritch, London, 521 pp.
Cretella, M. (2010) The complete collation and dating of the section Zoologie of the Coquille voyage. Bollethino
Malacologico, 4, 83–103.
Cory, C.B. (1918) Catalogue of Birds of the Americas. Field Museum Natural History Publications, Zoological Series, 13 (Part
2), No. 1, 1–315.
Coues, E. 1979. Second Instalment of American Ornithological Bibliography. Bulletin of the United States Geological and
Geographical Survey of the Territories, 5 (2), 239–696.
DeLattre, P.A. (1843) Oiseaux-mouches nouveaux au peu connus, découverts au Guatimala. L’Echo du Monde Savante, Series
2, 7 (No. 45), cols. 1068–1070.
DeLattre, P.A. & Bourcier, J. (1846) Description de quinze espèces nouvelles, faisant partie des collections rapportées par M.
Ad. De Lattre. Revue Zoologique, 1846, 305–312.
DeLattre, A. & Lesson, R.P. (1839) Oiseaux-mouches nouvelles ou très rares, découverts par M. De Lattre dans sou voyage à
Amérique et décrits par MM. De Lattre et Lesson. Revue Zoologique, 1839, 13–20.
Deppe, W. (1830) Preis-Verzeichnise der Säuethiere, Vögel, Amphibien, Fische und Krabbe, welchen von den Herren Deppe
und Scheide in Mexico gesammelt worden. The author, Berlin, 7 pp.
Dickinson, E.C. & Remsen, J.V. Jr. (Eds.) (2013) The Howard and Moore Complete Checklist of the Birds of the World. Vol. 1.
Non-passerines. Aves Press, Ltd., Eastbourne, 461 pp.
Dickinson, E.C., Overstreet, L.K., Dowsett, R.J. & Bruce, M.D. (2011) Priority! The dating of scientific names in ornithology:
a directory to the literature and its reviewers. Aves Press, Ltd., Northampton, 319 pp.
Elliot, D.G. (1871) Descriptions of two new species of humming-birds belonging to the genera Eupherusa and Cyanomyia.
Annals and Magazine of Natural History, Series 4, 2 (No. 66), 267–268.
Elliot, D.G. (1879) Classification and Synopsis of the Trochilidae. Smithsonian Contributions to Knowledge, No. 317, 1–277.
Friedmann, H., Griscom, L. & Moore, R.T. (1950) A Distributional Checklist of the Birds of Mexico. Part 1. Pacific Coast
Avifauna No. 29. Cooper Ornithological Club, Berkeley, California, 202 pp.
García-Deras, G., Cortés-Rodríguez, N., Honey, M., Navarro-Sigüenza, A.G., García-Moreno, J. & Hernández-Baños, B.E.
(2008) Phylogenetic relationships within the genus Cynanthus (Aves: Trochilidae), with emphasis on C. doubledayi.
Zootaxa, 1742, 61−68.
Gervais, P. (1835) Oiseaux-mouches. Ornismya Lesson. Magasin de Zoologie, Classe II, 1835, 46–49, pl. 43.
Gmelin, J.F. (1788) Caroli Linné Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordenes, Genera, Species, cum
Caracteribus, Differentiis, Synonymis, Locis. Tomus 1. Edicion Decimotertia, auct, reformata. Beer, Leipzig, 500 pp.
González, C., Ornelas, J.F. & Gutiérrez-Rodríguez, C. (2011) Selection and geographic isolation influence hummingbird
speciation: genetic, acoustical and morphological divergence in the Wedge-tailed Sabrewing (Campylopterus excellens).
Evolutionary Biology, 2011, 11–38.

STILES ET AL.
414
·
Zootaxa 4353 (3) © 2017 Magnolia Press
https://doi.org/10.1186/1471-2148-11-38
Gould, J. (1847) Drafts for an arrangement of the Trochilidae, with descriptions of some new species. Proceedings of the
Zoological Society of London, 15, 94–96.
Gould, J. (1848) Drafts for an arrangement of the Trochilidae, continued from part 15. Proceedings of the Zoological Society of
London, 16, 11–14.
Gould, J. (1850) On six new species of humming-birds. Proceedings of the Zoological Society of London, 18, 162–166.
Gould, J. (1851) On some new species of Trochilidae. Proceedings of the Zoological Society of London, 19, 115–116.
https://doi.org/10.1111/j.1096-3642.1851.tb01139.x
Gould, J. (1852) Monograph of the Trochilidae. Part 4. The author, London, pl. 12 & text.
Gould, J. (1853a) Description of five new species of humming birds. Proceedings of the Zoological Society of London, 21, 61–
62.
https://doi.org/10.1111/j.1469-7998.1853.tb07183.x
Gould, J. (1853b) Monograph of the Trochilidae. Part 5. The author, London, pl. 14 & text.
Gould, J. (1853c) Monograph of the Trochilidae. Part 6. The author, London, pl. 14 & text.
Gould, J. (1857) Monograph of the Trochilidae. Part 14. The author, London, pl. 12 & text.
Gould, J. (1858) Monograph of the Trochilidae. Part 16. The author, London, pl. 12 & text.
Gould, J. (1859) Description of four new species of humming-birds from Mexico. Annals and Magazine of Natural History,
Series 3, 4, 96–98.
Gould, J. (1861) Introduction to the Trochilidae. 8
th
Edition. Taylor & Francis, London, 216 pp.
Gould, J. (1862) Exhibition and description of two new species of humming-birds from Ecuador. Proceedings of the Zoological
Society of London, 1862, 124–125.
Gould, J. (1866) Leucippus chlorocercus. In: Sclater, P.L. & Salvin, O., Catalogue of birds collected by Mr. E. Bartlett on the
River Ucayali, eastern Peru, with notes and descriptions of new species. Proceedings of the Zoological Society of London,
1866, pp. 194.
Gray, G.R. (1840) A List of the Genera of Birds, with an Indication of the Typical Species of each Genus. R. & J.E. Taylor,
London, 115 pp.
Gray, G.R. (1855) Catalogue of the Genera and Subgenera of Birds contained in the British Museum. Trustees of the British
Museum, London, 192 pp.
Heine, F. (1863) Trochilidica. Journal für Ornithologie, 11, 172–217.
https://doi.org/10.1007/BF02004901
Hellmayr, C.E. (1913) Critical notes on the types of some little-known species of Neotropical birds, part 2. Novitates
Zoologicae, 20, 251–254.
Hernández, B.E., Zamudio-Beltrán, L.E., Eguiarte-Fruns, L.E., Klicka, J. & García-Moreno, J. (2014) The Basilinna genus
(Aves: Trochilidae): an evaluation based on molecular evidence and implications for the genus Hylocharis. Revista
Mexicana de Biodiversidad, 85, 797−807.
https://doi.org/10.7550/rmb.35769
Hilty, S.L. (2002) A Guide to the Birds of Venezuela. Princeton University Press, Princeton, New Jersey, 878 pp.
Hilty, S.L. & Brown, W.L. (1986) A Guide to the Birds of Colombia. Princeton University Press, Princeton, New Jersey, 836 pp.
Howell, S.N.G. (1993) Taxonomy and distribution of the hummingbird genus Chlorostilbon in Mexico and northern Central
America. Euphonia, 2, 25−37.
Howell, S.N.G. & Webb, S. (1995) A Guide to the Birds of Mexico and Northern Central America. Oxford University Press,
Oxford, 851 pp.
International Commission on Zoological Nomenclature (ICZN) (1999) International Code of Zoological Nomenclature. 4
th
Edition. International Trust for Zoological Nomenclature, London, 306 pp.
Jiménez, A.R. & Ornelas, J.F. (2016) Historical and current introgression in a Mesoamerican hummingbird species complex: a
biogeographic perspective. PeerJ, 4, e556.
https://doi.org/10.7717/peerj.1556
Krabbe, N. & Ridgely, R.S. (2010) A new subspecies of the Amazilia hummingbird from southern Ecuador. Bulletin of the
British Ornithologists’ Club, 130, 3–7.
Lacépède, M. Comte de (1799) Tableau des Sous-classes, Divisions, Ordres et Genres des Oiseaux. Libraire Plassens, Paris, 20
pp.
Lawrence, G.N. (1855) Description of a new species of humming bird of the genus Mellisuga Brisson, with a note on Trochilus
aquila Bourcier. Annals of the Lycaeum of Natural History of New York, 6, 137–142.
https://doi.org/10.1111/j.1749-6632.1858.tb00354.x
Lesson, R.P. (1826–1839) Voyage autour du Monde, Enterprise par Ordre du Gouvernement sur la Corvette La Coquille,
Zoologie. Vol. 1 & 15. Pourrat Frères, Paris, 683 pp. (1830) & 335 pp. (1830).
Lesson, R.P. (1829–1830) Histoire Naturelle des Oiseaux-Mouches, Arthus Bertrand, Paris, 330 pp.
Lesson, R.P. (1831–1832a) Histoire Naturelle des Colibris, suive d’un Supplément à l’Histoire Naturelle des Oiseaux-Mouches.
Arthus Bertrand, Paris, 196 pp.
Lesson, R. P. (1832b-1833) Les Trochilidés, ou les Trochilidés et le Oiseaux-Mouches, suivez d’un Index Générale, dans sont
décrits et Classées méthodiquement toutes les Races et Espèces du Genre Trochilus. Arthus Bertrand, Paris, 171 pp, plus

Zootaxa 4353 (3) © 2017 Magnolia Press
·
415
A NEW CLASSIFICATION OF THE TROCHI LINI
48 pp. (Index, numbered separately).
Lesson, R.P. (1843) L'Echo du Monde Savante, Series 2, 7 (No. 32), cols.756–757.
Lesson, R.P. & Garnot, P. (1827) Voyage autour du Monde, Enterprise par Ordre du Gouvernement sur la Corvette La
Coquille, Zoologie, 4, pl. 31, fig. 3.
Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, Secundum Classis, Ordenis, Genera, Species, cum
Caracteribus, Differentiis, Synonymis, Locis. Tomus 1. Edicion Decimal reformata. Impensis Direct. Laurentii Salvii,
Holmiae, 824 pp.
Linnaeus, C. (1766) Systema Naturae per Regna Tria Naturae, Secundum Classis, Ordenis, Genera, Species, cum
Caracteribus, Differentiis, Synonymis, Locis. Tomus 1. Edicion Duodecima reformata. Impensis Direct. Laurentii Salvii,
Holmiae, 865 pp.
Lozano-Jaramillo, M., Rico-Guevara, A. & Cadena, C.D. (2014) Genetic differentiation, niche divergence and the origin and
maintenance of disjunct distribution in the Blossomcrown, Anthocephala floriceps (Trochilidae). PLoS ONE, 9, e108345,
1–11.
McGuire, J.A., Witt, C.C., Remsen, J.V. Jr., Corl, A., Rabosky, D.L., Altshuler, D.L. & Dudley, R. (2014) Molecular
phylogenetics and the diversification of hummingbirds. Current Biology, 24, 1–7.
https://doi.org/10.1016/j.cub.2014.03.016
Meyer de Schauensee, R. (1966) The Species of Birds of South America and their Distribution. Livingston Publishing Co.,
Philadelphia, Pennsylvania, 577 pp.
Mulsant, M.E. & Verreaux, E. (1874) Histoire Naturelle des Oiseaux-mouches ou Colibris, constituent la Famille des
Trochilidés. Vol. 1. Bureau de la Société Linnéenne, Lyon, France, 405 pp.
Mulsant, M.E. & Verreaux, E. (1877) Histoire Naturelle des Oiseaux-mouches ou Colibris, constituent la Famille des
Trochilidés. Vol. 4. Bureau de la Société Linnéenne, Lyon, 353 pp.
Mulsant, M.E, Verreaux, J. & Verreaux, E. (1866) Essai d’un Classification des Trochilidés ou Oiseaux-Mouches. Mémoires de
la Société Impériale des Sciences Naturelles de Cherbourg, Series 2, 7, 140–252.
Nelson, E.W. (1911) Description of a new genus and species of hummingbird from Panama. Smithsonian Miscellaneous
Collections, 56 (No. 21), 1–2.
Nelson, E.W. (1912) Descriptions of new genera, species and subspecies of birds from Panama, Colombia and Ecuador.
Smithsonian Miscellaneous Collections, 60 (No. 3), 6–8.
Özdikmen, H. (2008) Neodamophila nom. nov., a replacement name for the bird genus Damophila Reichenbach, 1854 (Aves:
Apodiformes: Trochilidae). Munich Entomological Journal, 3, 171–173.
Peters, J.L. (1945) Check?list of Birds of the World. Vol. 5. Harvard University Press, Cambridge, Massachusetts, 306 pp.
Reich, G.C. (1795) Kürze Beschreibung verschiedener neuen, oder noch wenig bekannten Thiere, welche Herr Le Blond für
naturforschenden Gesellschaft zu Paris aus Cayenne Geschenk Übersicht bat. Magazin des Thierreich, 1795 (3), 128–134.
Reichenbach, L. (1854) Aufzählung der Colibris oder Trochilideen in ihrer wahrenatürlichen Verwandtschaft, nebst Schlüssel
ihrer Synonymik. Journal für Ornithologie, 1853 (Extraheft), 1–24.
Reichenbach, L. (1855) Trochilidarum Enumeratio. Cabanisii Diario Ornitologico, 2, 1–12.
Remsen, J.V. Jr., Stiles, F.G. & McGuire, J.A. (2015) Classification of the Polytminae (Aves: Trochilidae). Zootaxa, 3957 (1),
143–150.
https://doi.org/10.11646/zootaxa.3957.1.13
Ridgway, R. (1911) The Birds of North and Middle America. Bulletin of the United States National Museum, No. 50 (Part 5),
1–859.
Rodríguez-Gómez, F. & Ornelas, J.F. (2015) At the passing gate: past introgression in the process of speciation between
Amazilia violiceps and A. viridifrons hummingbirds along the Mexican Transition Zone. Journal of Biogeography, 47,
1305–1318.
https://doi.org/10.1111/jbi.12506
Salvin, O. (1892) Suborder Trochili. In: Catalogue of the Birds in the British Museum. Vol. 16. Trustees of the British Museum,
London, pp. 27–433.
Schuchmann, K.-L. (1999) Family Trochilidae (hummingbirds). In: del Hoyo, J., Elliot, A. & Sargatal, J. (Eds.), Handbook of
the Birds of the World. Vol. 5. Barn-owls to Hummingbirds. Lynx Edicions, Barcelona, pp. 468–680
Schulenberg, T. S., Stotz, D. F., Lane, D. F., O’Neill, J. P. & Parker, T. A., III. Birds of Peru. Princeton University Press,
Princeton, New Jersey, 656 pp.
Sclater, P.L. (1879) Thaumasius taczanowskii, sp. nov. Proceedings of the Zoological Society of London, 1879, 146.
Simon, E. (1897) Catalogue des espèces actuellement connues de la famille des Trochilidés. L. Mulo, Paris, 416 pp.
https://doi.org/10.5962/bhl.title.137190
Simon, E. (1901) Descriptions de trois espèces de la famille de Trochilidae. Ornis, 11, 201–203.
Simon, E. (1910) Catalogue général des Trochilidés dans la République d’Écuador. Revue Française d’Ornithologie, 1 (17–
18), 257–270.
Simon, E. (1921) Histoire Naturelle des Trochilidae: Synopsis et Catalogue. Enciclopedia Roret, L. Mulo, Paris, 420 pp.
https://doi.org/10.5962/bhl.title.15230
Simon, E. & Hellmayr, C.E. (1908) Notes critiques sur quelques Trochilidae. Novitates Zoologicae, 15, 1–12.
Stiles, F.G., Piacentini, V. de Q. & Remsen, J.V. Jr. (2017) A brief history of the generic classification of the Trochilini (Aves :

STILES ET AL.
416
·
Zootaxa 4353 (3) © 2017 Magnolia Press
Trochilidae): the chaos of the past and problems to be resolved. Zootaxa, 4269 (3), 396–412.
https://doi.org/10.11646/zootaxa.4269.3.4
Stiles, F.G. & Skutch, A.F. (1989) A guide to the birds of Costa Rica. Cornell University Press, Ithaca, NY, 510 pp.
Stone, W. (1907) Changes in generic names of North American birds. The Auk, 24, 189–199.
https://doi.org/10.2307/4070852
Stone, W. (1918) Birds of the Panama Canal Zone, with special reference to a collection made by Mr. Lindsey L. Jewel.
Proceedings of the Academy of Natural Sciences of Philadelphia, 70, 239–280.
Swainson, W. (1827a) Synopsis of the birds discovered in Mexico by W. Bullock, F.L.S. and H.S., and Mr. William Bullock,
jun. Philosophical Magazine, New Series, 1, 433–442.
Swainson, W. (1827b) On several groups and forms in ornithology, not hitherto defined. The Zoological Journal, 3 (11), 343–
365.
Tschudi, J.J. von (1846) Untersuchung über die Fauna Peruanum. Ornithologie. Schleitlin und Zollikofer, Vienna, 75 pp. [pp.
242–316]
Vieillot, L.P. (1818) Oiseaux-Mouches. Tome 23. In: Nouveau Dictionnaire d’Histoire Naturelle. Imprimerie d’Abel Lanoe,
Paris, pp. 423–431.
Vieillot, L.P. (1823) Ornithologie. In: Bonnaterre, A. de & Vieillot, L.P., Tableau Encyclopédique et méthodique des Trois
Règnes de la Nature. Part 2. Imprimerie Agasse, Paris, pp. 1–1460.
Weller, A.A. (1999) On types of trochilids in the Natural History Museum, Tring II. Bulletin of the British Ornithologists’ Club,
119, 97–102.
Weller, A.-A. (2000) Biogeography, geographic variation, and taxonomy of the Neotropical hummingbird genus Polyerata
Heine, 1863 (Aves: Trochilidae). Bonner Zoologische Monographien, 46, 47–54.
Zimmer, J.T. (1950a) Studies of Peruvian birds, no. 58: the genera Chlorostilbon, Thalurania, Hylocharis and Chrysuronia.
American Museum Novitates, 1473, 1–27.
Zimmer, J.T. (1950b) Studies of Peruvian birds, no. 59: the genera Polytmus, Leucippus and Amazilia. American Museum
Novitates, 1475, 1–31.
Zoonomen (2009) Zoological Nomenclature Resource; the Richmond index: genera. Bird Division, National Museum of
Natural History, Washington, D. C. Online version 2009, by A. P. Peterson. Available from: http://www.zoonomen.net
(accessed 17 October 2017)

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APPENDIX 1. Genus-level names for the emeralds accepted and names placed in synonymy in this study. Citations to the literature are given here in abbreviated form, with pertinent page numbers for
each name, but full citations for the works containing all genus and type species names are given in the text. For original designations authors, dates and works of type fixation are as for the genus.
Genus Name1
Author, date, work
Type species
Type fixation
This study
Abeillia
Bonaparte, 1850a, Consp. Gen. Av., 1,
79
Ornismya abeillei Lesson & De Lattre, Rev. Zool., 1839, 16
Orig. desig. (monotypy)
Accepted name
Agyrtria
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus thaumantias Linnaeus, Syst. Nat., ed. 12, 1766, 190
Automatic desig. (replacement
name)
= Polytmus
Agyrtrina
Chubb, 1916, Bds. Brit. Guiana, I, 393
Trochilus whitelyi Boucard, 1893, Hum. Bd., 8
Orig. desig. (monotypy)
= Chrysuronia
Amazilia
Lesson, 1843, L'Echo Mond. Sav., ser.
2, 7, no. 32, col. 757
Ornismya rutila De Lattre, 1843, L'Echo Mond. Sav., ser. 2, 7, no. 45, col.
1069
Stone 1918, P.N. A. S. P., 70, 256
(subs. desig.)
Accepted name
Amazilis
Gray, 1855, Cat. Gen. Bds. Brit. Mus.,
23
Orthorhynchus amazilia (as amazilis) Lesson & Garnot, 1827, Voy. Autour
Monde, 4, pl. 31, fig. 3
Orig. desig. (monotypy)
Accepted name
Amizilis
Gray, 1840, List Gen. Bds., 14
Orthorhynchus amazilia (as amizili) Lesson & Garnot, 1827, Voy. Autour
Monde, 4, pl. 31, fig. 3 = latirostris Swainson, 1827a, Philos. Mag., 11, 441
Orig. desig. (monotypy)
=Cynanthus
Anthocephala
Cabanis & Heine, 1860, Mus. Hein., 3,
8
Trochilus floriceps Gould, 1853b, P. Z. S. L., 1853, 62
Orig. desig. (monotypy)
Accepted name
Aphantochroa
Gould, 1853c, Mon. Troch., 6, pl. 12
Trochilus cirrochloris Vieillot, 1818, Nouv. Dict., 23, 430
Orig. desig. (monotypy)
Accepted name
Arenella
Simon, 1921, Synops. Cat.
Trochilidae
, 331
Arinia boucardi Mulsant & E. Verreaux, 1877, Hist. Nat. Col. O-m., vol. 4,
194
Orig. desig. (monotypy)
repl. Arena, Arinia
(preocc.); unresolved
Basilinna
Boie, 1831, Isis v. Oken, 24(4), -col.
546
Trochilus leucotis Vieillot, 1818, Nouv. Dict., 23, 428
Gray 1855, Cat. Gen. Brit. Mus., 23
(subs. desig.)
Accepted name
Brabournea
Chubb, 1916, Bds. Brit. Guiana, 1,
394
Thaumasius taczanowskii Sclater, 1879, P. Z. S. L., 1879, 146
Orig. desig. (monotypy)
= Thaumasius
Campylopterus
Swainson, 1827b, Zool. J., 3, 58
Trochilus largipennis Boddaert, 1783, Pl. Enl. Hist. Nat., 41
Gray 1840, List Gen. Birds, 13
(subs. desig.)
Accepted name
Chalybura
(Agyrtria δ)
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus buffonii Lesson, 1832a, Hist. Nat. Col., 31
Elliot 1879, Class. Syn. Troch., 45
(subs. desig.)
Accepted name
Chionomesa
Simon, 1921, Synops. Cat.
Trochilidae, 320
Ornismya lactea Lesson, 1833, Hist. Nat. Col., 99
Present study (subs. desig.)
Accepted name
Chrysuronia
Bonaparte, 1850a, Consp. Gen. Av., 1,
75
Ornismya oenone Lesson, 1832a, Hist. Nat. Col., 157
Gray 1855, Cat. Gen. Brit. Mus., 23
(subs. desig.)
Accepted name
Cynanthus
Swainson, 1827a, Philos. Mag., 11,
441
Cynanthus latirostris Swainson, 1827a, 441
Stone 1905 Auk, 24, 192 (subs.
desig.)
Accepted name
Cyanochloris
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus coeruleogaster Gould, 1847, P. Z. S. L, 1847, 95
Gray 1855, Cat. Gen. Brit. Mus.,
142 (subs. desig.)
= Chalybura
Cyanomyia
Bonaparte, 1854, Rev. Mag. Zool., 253
Ornismya cyanocephala Lesson, 1829, Hist. Nat. Ois-Mouch., 45
Elliot 1879, Class. Syn. Troch., 197
(subs. desig.)
= Saucerottia
Cyanophaia
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus bicolor Gmelin, 1788, Syst. Nat., ed. 13, 496
Gray 1855, Cat. Gen. Brit. Mus.,
142 (subs. desig.)
= Riccordia
……continued on the next page

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APPENDIX 1. (Continued)
Genus Name1
Author, date, work
Type species
Type fixation
This study
Chlorestes
Reichenbach, 1854, J. f. Orn., 1, 7
Trochilus notatus Reich, 1795, Mag. Thierreich, 129
Salvin 1892, Cat. Birds Brit. Mus.,
16, 239 (subs. desig.)
Accepted name
Chlorostilbon
Gould, 1853a, Monogr. Troch., pt. 5,
pl. 14
Ornismya prasinus Lesson, 1830, Hist. Nat. Ois.-Mouch., 188 (=Trochilus
mellisugus Linnaeus, 1758, Syst. Nat., ed. 10, 121)
Zimmer 1950a, Am. Mus. Nov., 1473,
4 (subs. desig.)
Accepted name
Damophila
(Coeligena β)
Reichenbach, 1854, J. f. Orn., 1, 7
Ornismya julie Bourcier, 1842, Rev. Zool., 1842,73
Elliot 1879, Class. Syn. Troch., 233
(subs. desig.)
= Chlorestes
Doleromya
Bonaparte, 1854, Rev. Mag. Zool., 6,
249
Trochilus fallax Bourcier, 1843b, Rev. Zool.,1843, 103
Orig. desig. (monotypy)
= Leucippus
Elvira
Mulsant, E. & J. & E. Verreaux, 1866,
Mem. Soc. Cherb., ser. 2, no. 7, 176
Thaumatias chionura Gould, 1850, P. Z. S. L., 1850 ,162
Orig. desig. (monotypy)
= Microchera
Elliotia
Stiles, Remsen & McGuire (this study)
Trochilus chionogaster Tschudi, 1846, p. 247, pl. 22
Orig. desig. (this study)
Accepted name
Eucephala
(Hylocharis δ)
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus grayi De Lattre & Bourcier, 1846, Rev. Zool., 1846, 307
Orig. desig. (monotypy)
= Chrysuronia
Eupherusa
Gould, 1857, Monogr. Troch., pt. 14,
pl. 12
Ornismya eximia De Lattre, 1843, L’Echo Mond. Sav., ser. 2, 7, no. 45, col.
1069
Orig. desig. (monotypy)
Accepted name
Eupetomena
Gould, 1852, Monogr. Troch., pt.4, pl.
12
Trochilus macrourus Gmelin, 1788, Syst. Nat., ed. 13,487
Orig. desig. (monotypy)
Accepted name
Goethalsia
Nelson, 1912, Smiths. Misc. Coll.,
60(3), 1
Goethalsia bella Nelson, 1912, 1
Orig. desig. (monotypy)
= Goldmania
Goldmania
Nelson, 1911, Smiths. Misc. Coll.,
56(21), 6
Goldmania violiceps Nelson, 1911, 6
Orig. desig. (monotypy)
Accepted name
Hylocharis
Boie, 1831, Isis v. Oken, 24(4), col.
546
Trochilus sapphirinus Gmelin, 1788, Syst. Nat., ed. 13, 245
Gray 1840, List Gen. Birds, 14 (subs.
desig.)
Accepted name
Juliamyia
Bonaparte, 1854, Rev. Zool., 6, 255
Ornismya julie Bourcier, 1842, Rev. Zool., 1842, 3, 73
Orig. desig. (monotypy)
= Chlorestes
Klais (Basilinna β)
Reichenbach, 1854, J. f. Orn., 1, 13
Trochilus guimeti Bourcier & Mulsant, 1843, Ann. Soc. Sci. Lyon, 6,38
Orig. desig. (monotypy)
Accepted name
Lepidopyga
(Agyrtria γ)
Reichenbach, 1855, Troch. Enum., 7
Trochilus goudoti Bourcier, 1843, Rev. Zool., 1843, 100
Ridgway 1911, Birds N Mid. Am.,
no. 50, pt. 5, 537 (subs. desig.)
= Chrysuronia
Leucippus
Bonaparte, 1850a, Consp. Gen. Av., 1,
73
Trochilus fallax Bourcier, 1843, Rev. Zool., 1843, 103
Gray 1855, Cat. Gen. Brit. Mus., 21
(subs. desig.)
Accepted name
Leucochloris
(Agyrtria α)
Reichenbach, 1854, J. f. Orn., 1, 10
Trochilus albicollis Vieillot, 1818, Nouv. Dict., 23, 426
Orig. desig. (monotypy)
Accepted name
Leucolia
Mulsant, E & J. & E. Verreaux, 1866,
Mem. Soc. Cherb., ser. 2, no. 7, 172
Cyanomyia viridifrons Elliot, 1871, Ann. Mag. Nat. Hist., 2, 267
Elliot 1879, Class. Syn. Troch., 195,
present study
Accepted name
Microchera
Gould, 1858, Monogr. Troch., pt. 16,
pl. 12
Mellisuga albocoronata Lawrence, 1855, Ann. Lyc. New York, 6, 137
Orig. desig. (monotypy)
Accepted name
Orthorhynchus
Lacépède, 1799, Tabl. Mam. Ois., 10
Trochilus cristatus Linnaeus, 1758, Syst. Nat., ed. 10, 192
Gray 1840, List Gen. Birds, 14 (subs.
desig.)
Accepted name
……continued on the next page

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419
A NEW CLASSIFICATION OF THE TROCHI LINI
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APPENDIX 2. Proposal for a revised classification of the genera of Trochilini
The linear sequence below is derived from the genetic tree of McGuire (2014) illustrated in Fig. 1, and follows the
standard convention of listing least-diverse branches of a clade first, except for superspecies complexes, in which case
the species are listed by the convention of geographic distribution from northwestern to southeastern. Our procedure for
generic rearrangements is explained in the text. Species marked with an asterisk were not sampled by McGuire et al.
(2014) and their placement in the species sequences is tentative or unresolved; those with a dagger are recently extinct
species. Type species of each genus recognized here are distinguished by the species epithet in bold face and cited by
author and date; generic names used by Schuchmann (1999) and Dickinson and Remsen (2013) but synonymized or
mentioned here are given under the respective genera below, with their type species, authors and dates. See the text and
Appendix 1 for details of generic descriptions and type species designations. This classification recognizes 110 species
in 35 genera, with two species currently unplaced due to lack of genetic samples. We have synonymized seven genera
previously recognized: Cyanophaia, Aphantochroa, Elvira, Goethalsia, Agyrtrina, Lepidopyga and Juliamyia;
recognized nine genera considered as synonyms or not mentioned in recent lists (Schuchmann 1999, Dickinson &
Remsen 2013): Riccordia, Phaeoptila, Pampa, Leucolia, Uranomitra, Thaumasius, Amazilis, Talaphorus and
Chionomesa; and named one new genus, Elliotia. The generic name Agyrtria is rejected as not applicable within the
Trochilini (see Stiles et. al., 2017).
Phaeoptila Gould 1861
Phaeoptila sordidus (Gould, 1859)
Riccordia Reichenbach, 1854.
Syn: Cyanophaia Reichenbach, 1854; type, Trochilus bicolor Gmelin,1788
Note: R. bracei is tentatively placed here and adjacent to R. ricordii, with which it had been previously considered
conspecific; see Graves & Olson (1987).
Riccordia ricordii (Gervais, 1835)
Riccordia bracei (Lawrence)*†
Riccordia elegans (Gould)*†
Riccordia swainsonii (Lesson)
Riccordia maugaeus (Audebert & Vieillot)
Riccordia bicolor (Gmelin)
Cynanthus Swainson, 1827a. Note: if considered separate species, osberti* and salvini* would follow forficatus in the
linear sequence.
Cynanthus latirostris Swainson, 1827a
Cynanthus doubledayi (Bourcier)
Cynanthus auriceps (Gould)*
Cynanthus forficatus (Ridgway)
Cynanthus canivetii (Lesson)
Chlorostilbon Gould, 1853a
Chlorostilbon mellisugus (Linnaeus, 1758)
Chlorostilbon olivaresi Stiles*
Chlorostilbon gibsoni (Fraser)*
Chlorostilbon lucidus (Shaw)
Chlorostilbon poortmani (Bourcier)
Chlorostilbon stenurus (Cabanis & Heine)*
Chlorostilbon alice (Bourcier & Mulsant)*
Chlorostilbon russatus (Salvin & Godman)*
Chlorostilbon assimilis Lawrence
Chlorostilbon melanorhynchus Gould

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A NEW CLASSIFICATION OF THE TROCHI LINI
Basilinna Boie, 1831
Basilinna leucotis (Vieillot, 1818)
Basilinna xantusii (Lawrence)
Pampa Reichenbach, 1854
Pampa curvipennis (Deppe, 1830)*
Pampa excellens Wetmore
Pampa rufus (Lesson)
Klais Reichenbach, 1854
Klais guimeti (Bourcier & Mulsant, 1843)
Abeillia Bonaparte, 1850a
Abeillia abeillei (De Lattre & Lesson, 1839)
Orthorhynchus Lacépède, 1799
Orthorhynchus cristatus (Linnaeus, 1758)
Stephanoxis, Simon 1897. Note: for treatment of S. loddigesii as a separate species, see Cavarzere et al. (2014); citation
in text.
Stephanoxis lalandi (Vieillot, 1818)
Stephanoxis loddigesii (Gould)*
Anthocephala Cabanis and Heine, 1860. Note: for treatment of A. berlepschi as a separate species, see Lozano-
Jaramillo et al. (2014); citation in text.
Anthocephala floriceps (Gould, 1853b)
Anthocephala berlepschi Salvin*
Campylopterus Swainson 1827b. Note: C. phainopeplus is included based on traditional classifications and
morphological similarities and is placed in the sequence following Schuchmann (1999).
Campylopterus largipennis (Boddaert, 1783)
Campylopterus hemileucurus (Deppe)
Campylopterus hyperythrus Cabanis
Campylopterus duidae Chapman
Campylopterus villaviscensio (Bourcier)
Campylopterus falcatus (Swainson)
Campylopterus phainopeplus Salvin & Godman*
Campylopterus ensipennis (Swainson)
Chalybura Reichenbach, 1854
Chalybura urochrysia (Gould)
Chalybura buffonii (Lesson, 1832a)
Thalurania Gould, 1848
Thalurania glaucopis (Gmelin)
Thalurania watertonii (Bourcier)
Thalurania colombica (Bourcier)
Thalurania furcata (Gmelin, 1788)

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Microchera Gould, 1858
Syn: Elvira Mulsant, J. & E. Verreaux, 1866; type Thaumatias chionura Gould, 1850
Microchera albocoronata (Lawrence, 1855)
Microchera cupreiceps (Lawrence)
Microchera chionura (Gould)
Goldmania, Nelson 1911
Syn: Goethalsia Nelson, 1912; type Goethalsia bella Nelson, 1912
Goldmania bella (Nelson)
Goldmania violiceps Nelson, 1911
Eupherusa Gould, 1857
Eupherusa ridgwayi (Nelson)
Eupherusa poliocerca Elliot
Eupherusa cyanophrys Rowley & Orr
Eupherusa eximia (DeLattre, 1843)
Eupherusa nigriventris Lawrence
Phaeochroa Gould, 1861
Phaeochroa cuvierii (DeLattre & Bourcier, 1846)
Leucippus Bonaparte, 1850a
Leucippus fallax (Bourcier, 1843)
Thaumasius P. L. Sclater, 1879
Thaumasius baeri (Simon)
Thaumasius taczanowskii P.L. Sclater, 1879
Taphrospilus Simon, 1910
Taphrospilus hypostictus (Gould, 1862)
Eupetomena Gould, 1852
Syn: Aphantochroa Gould, 1853c; type Trochilus cirrochloris Vieillot, 1818
Eupetomena macroura (Gmelin, 1788)
Eupetomena cirrochloris (Vieillot)
Talaphorus Mulsant & E. Verreaux, 1874
Talaphorus chlorocercus (Gould, 1866)
Trochilus Linnaeus, 1758
Trochilus polytmus Linnaeus, 1758
Trochilus scitulus Brewster & Bangs
Leucolia Mulsant, J. & E. Verreaux 1866
Leucolia violiceps (Gould)

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A NEW CLASSIFICATION OF THE TROCHI LINI
Leucolia wagneri Phillips
Leucolia viridifrons (Elliot, 1871)
Saucerottia Bonaparte, 1850b. Note: See text for treatment of S. hoffmanni as a separate species from S. saucerottei.
The species cyanifrons is tentatively included adjacent to S. saucerottei because they have been treated as
forming a superspecies (Schuchmann 1999).
Saucerottia cyanocephala (Lesson)
Saucerottia beryllina (Deppe)
Saucerottia cyanura (Gould)
Saucerottia hoffmanni (Cabanis & Heine)
Saucerottia edward (DeLattre & Bourcier)
Saucerottia saucerottei (DeLattre & Bourcier, 1846)
Saucerottia cyanifrons (Bourcier)
Saucerottia castaneiventris (Gould)
Saucerottia viridigaster (Bourcier)
Saucerottia tobaci (Gmelin)
Amazilia Lesson, 1843
Amazilia rutila (De Lattre, 1843)
Amazilia yucatanensis (Cabot)
Amazilia tzacatl (De la Llave)
Amazilis Gray, 1855
Amazilis amazilia (Lesson & Garnot, 1827)
Uranomitra Reichenbach, 1854
Uranomitra franciae (Bourcier & Mulsant, 1846)
Chrysuronia Bonaparte, 1850a
Syn: Agyrtrina Chubb, 1916; type Trochilus whitelyi Boucard, 1893
Syn: Lepidopyga Bonaparte, 1850a; type: Trochilus goudoti Bourcier, 1843
Syn: Eucephala Reichenbach, 1854; type, Trochilus grayi Bourcier & Mulsant, 1846
Note: The taxon lilliae is tentatively placed here and adjacent to C. coeruleogularis based on assumed close relationship
to that species; see Schuchmann (1999), but recognition as a separate species from coeruleogularis is
controversial.
Chrysuronia versicolor (Vieillot)
Chrysuronia goudoti (Bourcier)
Chrysuronia oenone (Lesson, 1832a)
Chrysuronia coeruleogularis (Gould)
Chrysuronia lilliae (Stone)*
Chrysuronia humboldtii (DeLattre & Bourcier)*
Chrysuronia grayi (Bourcier & Mulsant)
Chrysuronia brevirostris (Lesson)
Chrysuronia leucogaster (Gmelin)
Leucochloris Reichenbach, 1854
Leucochloris albicollis (Vieillot, 1818)
Chionomesa Simon 1921
Chionomesa fimbriata (Gmelin)
Chionomesa lactea (Lesson, 1833)

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Zootaxa 4353 (3) © 2017 Magnolia Press
Hylocharis Boie, 1831
Hylocharis sapphirina (Gmelin, 1788)
Hylocharis chrysura (Shaw)
Elliotia Stiles, Remsen & McGuire, 2017
Elliotia chionogaster (von Tschudi, 1846)
Elliotia viridicauda (von Berlepsch)
Polyerata Heine, 1863
Polyerata rosenbergi Boucard
Polyerata amabilis (Gould, 1851)
Polyerata decora Salvin
Chlorestes Reichenbach, 1854
Syn: Juliamyia Bonaparte, 1854; type Ornismya julie Bourcier, 1842
Chlorestes candida (Bourcier & Mulsant)
Chlorestes eliciae (Bourcier & Mulsant)
Chlorestes cyanus (Vieillot)
Chlorestes julie (Bourcier)
Chlorestes notata (Reich, 1795)
Incertae sedis
“Amazilia” luciae (Lawrence)*
“Amazilia” boucardi (Mulsant)*
These two unsampled members of the amazilian complex are unplaced here: for details, see text.