Stephanoxis loddigesii (Gould, 1831)

Cavarzere, Vagner, Silveira, Luís Fábio, Vasconcelos, Marcelo Ferreira de, Grantsau, Rolf & Straube, Fernando Costa, 2014, Taxonomy and biogeography of Stephanoxis Simon, 1897 (Aves: Trochilidae), Papéis Avulsos de Zoologia (São Paulo) 54 (7), pp. 69-79 : 73-75

Stephanoxis loddigesii (Gould, 1831)

English name: Purple-crested Plovercrest.

Portuguese name: beija-flor-de-topete-roxo.

Trochilus Loddigesii GOULD in Vigors, 1831: 12. Rio Grande.

Cephalepis loddigesi Bonaparte, 1850: 83 .

Cephalepis loddigesii: REICHENBACH, 1854: 12 .

Orthorhynchus loddigesii Reichenbach, 1855: 9 . Orthorhynchus loddigesi: BURMEISTER, 1856: 352 .

Cephalolepis loddigesi Cabanis & Heine, 1860: 61 . [Emendation of Cephallepis Loddiges ].

Bellona loddigesi Verreaux & Verreaux, 1866: 75 .

Cephallepis loddigesi Mulsant & Verreaux, 1874 -77: 291.

Stephanoxis Loddigesi Simon, 1897: 40 . [New name for Cephallepis Loddiges and Cephalolepis Cabanis & Heine , both pre-ocupied].

Stephanoxis loddigesi: HARTERT, 1900: 214 .

Stephanoxis lalandi loddigesii [subspecies]: PETERS, 1945: 31.

Cephaloepis apirati BERTONI & BERTONI in Bertoni, 1901: 55.

Diagnosis: Males diagnosed by a violet-blue crest (1), brownish auriculars (2), chin and throat (3), blackish violet-blue abdomen (4) and, as in females, yellowish green back (5) and crown (6) ( Table 1, Fig. 1 ).

Nomenclature

Vigors (1831) described Trochilus Loddigesii Gould , while acknowledging that the name was dedicated to Loddiges by J. Gould, to whose collection the type specimen belonged. According to article 50.1.1 of the International Code of Zoological Nomenclature ( ICZN, 1999), hereafter the Code, “…if it is clear from the contents that some person other than an author of the work is alone responsible both for the name or act and for satisfying the criteria of availability other than actual publication, then that other person is the author of the name or act.” This statement deems Stephanoxis loddigesii (GOULD in Vigors, 1831) both available and valid, satisfying the criteria of Articles 8 (what constitutes published work) and 10 – 12 (provisions and requirements conferring availability) of the Code. In the same publication, Loddiges (1831: 12) states that this “…species belonged to a genus which he [himself] had distinguished among the Trochilidae by the name of Cephallepis …” However, this new genus-group name was pre-occupied, leading Simon (1897) to provide a substitute name (nomen novum) for the genus (see also Cory, 1918; Peters, 1945). Namely, Cephallepis was already in use ( Rafinesque & Gill, 1810) while Cephalolepis was an attempt by Cabanis & Heine to amend Loddiges’s genus; Cabanis & Heine, however, failed to realize its previous applications ( Dumeril & Bibron, 1844; Agassiz, 1847). It is unclear to us why Peters (1945) considered Cephaloepis apirati BERTONI & BERTONI in Bertoni (1901) an emendation, since the author explicitly mentioned the etymology of this genus was unknown. He did consider, however, that Cephaloepis apirati could be a variation of Vieillot’s Trochilus Delandi (sic). It is, in fact, a synonym of Stephanoxis loddigesii .

Biogeography

The Serra do Mar has long been known to be an area of endemism for Atlantic forest vertebrates (e.g., Müller, 1973; Kinsey, 1982; Cracraft, 1985; Stattersfield et al., 1998; Costa et al., 2000), including passerine birds ( Silva et al., 2004). Müller was the first to propose a “Paulista” centre, suggesting that the vertebrate fauna in northern São Paulo is more closely related to the northern Atlantic forest than to the southern Serra de Paranapiacaba. This was corroborated by Costa et al. (2000), who came to the same conclusion, using a parsimony analysis of endemism with primates and small mammals, even though their grids were arbitrarily defined at a rough scale of 275 × 275 km. Nevertheless, this pattern was not recovered by Silva et al. (2004), who used the same biogeographic method to identify areas of ende- mism for passerine birds with 1 × 1 degree quadrats (ca. 110 km 2). The latter authors concluded there are four areas of endemism within the Atlantic forest, one consisting of the south-eastern portion of the domain, from the states of Espírito Santo south to northern Santa Catarina, which includes the Serra do Mar and Serra de Paranapiacaba in a single area.

Recently, the use of ecological niche models under paleoclimates and phylogeographic analyses have allowed the re-evaluation of some traditional hypotheses based on empirical species ranges and past and current disposition of forests and terrains. This includes the role of forest refugia in the diversification of Neotropical birds (e.g., Haffer 1967, 1969). Carnaval & Moritz (2008), modelling the spatial range of the coastal Brazilian Atlantic forest under different climatic scenarios, suggested the existence of stable forest refugia north and south of the São Francisco River, namely the Pernambuco and Bahia refuges, respectively. The high levels of endemism south of the Doce River could not, however, be explained as this region was predicted to bear less historical stability.

Despite the existence of two Plovercrest species, there are at least three other pairs of montane sister taxa with similar distributions in south-eastern Brazil, i.e. populations centred in northeast and south-western São Paulo segregated by a kilometric gap between the Serra do Mar (just north of the Baixada Santista) and Serra de Paranapiacaba. These include the Mousecoloured Tapaculo Scytalopus speluncae / Scytalopus sp. nov. ( Mata et al., 2009), the Buff-throated Poospiza lateralis / Gray-throated Warbling-Finch Poospiza cabanisi ( Assis et al., 2007) , and the Brown-breasted Pygmy-Tyrant Hemitriccus o. obsoletus / Hemitriccus o. zimmeri ( Straube & Di Giácomo, 2007: 61). There are also at least three species that have the same distribution pattern but that show no apparent break in plumage or voice consistency. These are the Graybellied Spinetail Synallaxis cinerascens, Serra do Mar Tyrannulet Phylloscartes difficilis and Great Pampa- Finch Embernagra platensis . Furthermore, Cabanne et al. (2007) revealed a cryptic genetic diversity within the Lesser Woodcreeper Xiphorhynchus f. fuscus , which has a large genetic distance observed in northern and southern São Paulo, exhibiting a range remarkably similar to Stephanoxis spp.

We followed the scenario in which Carnaval & Moritz’s (2008) and Carnaval et al. (2009) proposed that historical refuges south of the Doce River had existed and were less stable, but not absent (see also Maldonado-Coelho, 2012). Silva et al. (2012) systematically tested in how many vertebrate components the Brazilian Atlantic Forest could be divided by analysing existing empirical data. They found a marked east-west differentiation on the southerly portion of this domain, which may be credited either to the uplift of the Serra do Mar and Serra da Mantiqueira, recent population expansions within those forests ( Cabanne et al., 2008; D’Horta et al., 2011) or local topography that suffered contrasting modifications across time, making the southern Atlantic forest of Brazil unsuitable for an exuberant forest on several occasions. Since the Atlantic forest is composed of a complex mosaic of vegetation and topography with potential for vertical migration ( Por, 1992), we suggest Stephanoxis , Scytalopus , Hemitriccus and Poospiza may have evolved in cool and humid refuges (perhaps in higher elevations) during glacial periods. Other ecologically plastic species which were able to survive in expanding and retracting forests may have occupied surrounding habitats, therefore remaining phenotypically stable due to free genetic flow. Only older, true highland lineages, which supposedly remained isolated in mountaintops, are obviously morphologically distinguished at present. The ancestral populations of both lineages of Stephanoxis may have expanded northward during cooler and humid events. This follows a pattern similar to that observed for the Paraná pine Araucaria angustifolia (see Hueck, 1953, 1972; Behling, 1998, 2002), several other bird taxa ( Vasconcelos & Rodrigues, 2010; Freitas et al., 2012), a rodent ( Gonçalves et al., 2007) and bees ( Silveira & Cure, 1993), although not necessarily in the same time cycles. It seems probable that during one (or more) warmer and drier cycles, a population retract- ed southward and another moved to lower latitudes, becoming trapped in these south-eastern highlands, leading to vicariance. Although the barrier between both species is located in São Paulo, it is possible that the real vicariant barrier occurred elsewhere and that subsequent population expansions led to the current gap between the Serra do Paranapiacaba/Serra do Mar. A similar pattern was recently recognized in eastern Brazil for another highland hummingbird genus ( Vasconcelos et al., 2012).

We predict several more species occurring in this region may show recent populational expansions between extreme latitudes in São Paulo. This viewpoint is in accordance with phylogeographies of X. f. fuscus and Pyriglena leucoptera , Atlantic forest endemic bird species found from sea level to montane forests ( Cabanne et al., 2008; Maldonado-Coelho, 2012). We strongly suggest more phylogeographic studies be developed within those areas, which could account for overlooked biogeographic patterns. Although unsurprising to recently witness the discovery of a new area of endemism on the under-studied north-western Brazilian Amazonia ( Borges & Silva, 2012), how exciting would it be to suggest a novel insight into the biogeography of the scientifically best known rain forest of Brazil?

Figures

FIGURE 1: Stephanoxis lalandi (left) and Stephanoxis loddigesii (right). Males above, females below. Plate by Rolf Grantsau.