Catasticta lycurgus ( Godman & Salvin, 1880 )

Huertas, Blanca, Prieto, Carlos, Montero, Fredy, Adams, Mike, Crom, Jean François Le, Bollino, Maurizio, Correa-Carmona, Yenny & Padrón, Pablo Sebastián, 2021, One Hundred Years of Solitude: rediscovery of Catasticta lycurgus (Godman & Salvin, 1880), a yellow butterfly from the Sierra Nevada de Santa Marta, Colombia (Lepidoptera: Pieridae), Zootaxa 4975 (1), pp. 176-186 : 178-184

Catasticta lycurgus ( Godman & Salvin, 1880)

Holotype, male ( Figs. 1 a–c )

Catasticta Butler, 1870 “ Euterpe lycurgus , sp. n. ” Godman & Salvin, 1880: 132, Pl. IV: Fig. 15.

Type locality: “on the road from San Sebastian to Atánquez” ( Godman & Salvin, 1880: 132). Holotype (by monotypy) (Plate 23: Figs 177, 178).

Additionally, to the labels described above, the holotype of C. lycurgus has the following labels as listed in Abadjiev (2006):

(1) printed (on white paper) “S. Sebast. to | Atanques [sic] | F. Simons”; (2) circle printed (on white paper with red frame) “ Type ”; (3) printed (on white paper) “Godman-Salvin. | Coll.B.M. 1909-28.”; (4) printed (on white paper) “Photographed by | B. D’Abrera 77/78”; (5) printed (on white paper) “BMNH(E) #662336”.

A modern code has been placed recently ( NHMUK 014172794), meaning the specimen has been digitally photographed and is available online in the NHM data portal www.data.nhm.ac.uk.

Other non-type specimens ( Figs. 2 a–b )

1 male: Colombia, Magdalena, M. Adams & G. Bernard. / E. Above San Pedro, 2,900 metres, 7. viii. 1972. 2011 / G1.71/72 A/B. / BMNH(E)1203312 /NHMUK010430776 ( NHMUK) .

2 males: Col., Magdalena , Sierra Nevada, San Pedro. 2,600–2,700 m, 10–12/01/2013, codes, b376, b377 ( RCCP) .

4 males: Colombia – Depto. Magdalena / San Pedro de la Sierra – SNSM / 10°53’52.85”N – 73°57’7.82”W / III.2016, 3,000–3,150m / Leg. F. Montero ( MBLI) .

9 males: San Pedro de la Sierra Nevada, Mgd. 3,100 m. i. 2013. Leg. F. Montero ( JFLC) .

Re-description of the male and illustration of its genitalia ( Fig. 2a–b, f )

FW with slightly rounded apex and outer margin indented at M2-M3; HW with smooth outer margin and rounded anal lobe.

Dorsal surface: Ground colour of both wings brown (PANTONE® 1545 C). FW with a complete series of yellow (PANTONE® 7404 C) discal, submarginal and marginal spots; marginal spots slightly elongated; discal spots in space 1b-2–3 extended inward to touch cellular vein, those in space 1a–1b extended inwardly nearly to base of wing. Cell with inner two thirds yellow from base to apex. HW with complete series of yellow discal and submarginal spots; marginal spots lighter in colour; submarginal spots chevron-shaped; marginal spots triangular.

Ventral surface: FW with a complete series of discal, submarginal and marginal spots; marginal spots elongated. Discal spots light yellow, those in space 1b-2–3 extended inward to touch cellular vein, and those in space 1a–1b extended inwardly nearly to base of wing. Cell with outer third light yellow, and an axial yellow streak in basal two thirds. HW with general pattern as in Catasticta incerta (Dognin, 1888) (see below).

Genitalia ( Fig. 2f ): little or no attention has been given by past authors to the morphology of male genitalia of the genus Catasticta for their scarce complexity and simple structure. Klots (1933) associates the genus Catasticta with the genera Eucheira Westwood , Neophasia Behrand , and Archonias Hübner because they are “ all characterized most strikingly by an extreme development in size of the tegumen, with a correlated decrease in size of the articulatory process and an extreme shortening of the uncus ”. Eitschberger & Racheli (1998) only hint at the simple and little differentiated structure of the genitals in the different species of the genus. Apart from these notations, Bollino & Costa (2007) “ find no evidence in the literature of any species-specific differences in the various taxa of the genus Catasticta ” except an unconfirmed difference between two species reported by Wojtusiak (1998).

However, all this does not exclude that future research may demonstrate the existence of species-specific, or more likely group-specific characters, in the morphology of male genitalia of Catasticta . For this reason we illustrate the apparatus of a recently captured male.

Description of the female ( Fig. 2c–d )

Dorsal surface: Ground colour of both wings deep brown (PANTONE® 439 C). FW with a complete series of pale yellow (PANTONE® 600 C) discal, submarginal and marginal spots; submarginal and marginal spots obscured by brown scales; marginal spots slightly elongated; discal spot in space 1b extended inward to touch cellular vein only near base of vein CU2, those in space 2–3 extended inward to touch cellular vein. Cell with outer yellow spot touching the cellular vein. HW with complete series of slightly more saturated yellow (PANTONE® 393 C) discal, submarginal and marginal spots; the submarginal and marginal spots obscured by brown scales; submarginal spots chevron-shaped; marginal spots triangular.

Ventral surface: FW with a complete series of discal, submarginal and marginal spots; marginal spots elongated. Discal spots light yellow, with same extension as on dorsal surface. HW with general pattern as in Catasticta incerta (Dognin, 1888) (see below).

1 female: Col., Magdalena , Sierra Nevada, San Pedro, 2,600 m, 11/01/2013, code: b375. 10°54’48”N – 73°57’51”W / ( RCCP) .

Distribution and biology

Although there was no record of the exact location of the type locality as the labels of the type specimen of Catasticta lycurgus do not hold other collecting data rather than ‘from S[an] Sebast[ian de Rabago] to Atanques [sic] [Santa Marta, Colombia]’ ( Fig. 1b ). Simons, independently, wrote a few articles in specialised geographical journals giving exhaustive accounts of his expeditions to Santa Marta (and Guajira), including detailed maps drawn by himself ( Simons 1879; Simons 1881; Gómez-Creutzberg 2017). He wrote: “The Indian track between San Sebastian and Atánquez is very heavy walking, owing to the steep ascents. Nearly all along the road are Indian huts and plantations, especially at Templado” (op.cit.). Simons presented in one of his best known maps of the Sierra Nevada de Santa Marta, a drawing of a horizontal section between San Sebastián and Atánquez including a vertical scale of the altitude ( Simons 1879). Godman and Salvin also included details of the collecting localities recorded by Simons in his paper as follows: ‘ San Sebastian is located on the south-western slope of the Sierra Nevada, with an elevation of 6,700 ft (c. 2,000 m) and Atánquez, is located in the south-eastern slope of the [Sierra] Nevada, six hours’ journey from Valle Dupar [sic] and an elevation of 2,800 ft (c. 853 m) ’ ( Godman & Salvin 1880). Despite the uncertainties about the exact location of the type’s locality of origin, we do not believe it necessary to restrict the type locality, however, using the above information and Simons’ map which includes coordinates ( Simons 1879) we made an estimate of the possible coordinates for the type locality ( Fig. 3a ).

This century, Catasticta lycurgus has been recorded in elevations from 2,600 m up to 3,150 m (see non-type specimens) only in habitats in the Sierra Nevada de Santa Marta in Colombia. This time, precise coordinates of the habitats in which this species is found have been taken ( Fig. 3a ). In January 2013 CP found a relatively large population at 2,700m on a ridge of the track leading from San Pedro to the Río Frío lagoons. It appears that the species is common in certain years since at least 20 relatively fresh individuals were registered flying in a short period of time. Two males and one female were collected along the way. Specimens were rarely observed ‘mud-puddling’ ( Fig. 3c–d ). Although authors FM and CP have visited the area for fieldwork on various occasions, there are no observation of adult specimens feeding on flowers nor information on immature stages. This species has been observed flying from around 0800 until 1400 hours. It is found most often near the ecotone between páramo and montane forest, usually near the border of the forest ( Fig. 3b ). Occasionally, some individuals were observed flying across open areas or flying in forest patches. Almost always, the males continue flying and will chase away other males of the same species from their perches. Adult males suddenly stop their wanderings and congregate over the foliage of prominent branches, a typical behaviour known as hill-topping.

Systematic relations among the genus Catasticta

Catasticta lycurgus was originally described as Euterpe lycurgus by Godman & Salvin (1880). Since then, it has not been studied further, presumably due to a lack of specimens. Therefore, its relationship inside Catasticta remains unclear. Talbot (1932) considered C. lycurgus as a subspecies of C. nimbice (Boisduval, 1836) probably due to the similarity of the dorsal wing pattern to C. n. bryson Godman & Salvin, 1889 . However, the scheme of the ventral wing pattern of C. lycurgus is completely different from that of C. n. bryson .

Reissinger (1972) included C. lycurgus in the subgenus Catasticta , and in the subgroup nimbice , which included C. nimbice from Mexico and C. ochracea (Bates, 1864) from Guatemala and C. bryson from Costa Rica, the latter ones being currently regarded as subspecies of C. nimbice ; C. peruviana Joicey & Talbot, 1918 from Peru (currently recognized as a subspecies C. prioneris peruviana but most probably a valid species by its own); C. reducta reducta Butler, 1896 from Ecuador; C. reducta microperuviana Reissinger, 1972 from Peru; C. reducta boliviana Butler, 1896 from Bolivia; C. butleria Brown & Gabriel 1939 (a valid subspecies, now generally treated as C. reducta butleria ) from Peru; C. seitzi Lathy & Rosenberg 1912 , from Colombia; C. grossana Brown & Gabriel 1939 (currently C. philone grossana ) from Colombia and C. sinapina Butler, 1896 from Peru.

Eitschberger & Racheli (1998), in their subdivision of Catasticta in groups and subgroups based on the pattern designs on the hindwing underside, considered C. lycurgus inside the “ aureomaculata ” group which included C. incerta , C. fulva Joicey & Rosenberg, 1915 and C. lycurgus , the last including two subspecies C. l. lanceolata Lathy & Rosenberger, 1912 and C. l. tolima Fassl, 1915 .

Bollino & Rodríguez (2003) discussed these taxonomical combinations and removed the two subspecies from C. lycurgus , so revalidating C. lanceolata as a full species, with C. tolima as its subjective synonym. Le Crom et al. (2004) diagnosed it and sketched out its distribution. Finally, Lamas (2004), recognised C. lycurgus as a valid monotypic species.

Despite what has been said, the systematic relationships between C. lycurgus and the other species of Catasticta are not yet clear. One of the criteria of classification established by Eitschberger & Racheli (1998) states that “The design [on the hindwing underside] is complex and it is unlikely that it may be sustained only by convergence”; for the same reason we could suppose that the similarity on the hindwing underside of two or more taxa cannot be supported by convergence, but indicates that, most likely, these taxa are phylogenetically related. Examining in detail the FW and HW morphology in C. lycurgus , the species is very similar to C. incerta . Females of both species are also very similar to each other. The chorology and altitudinal distribution of both species would also suggest that possibly a common ancestor is shared: C. lycurgus is found at Sierra Nevada de Santa Marta between 2,600 to 3,150 m and C. incerta is distributed from the Colombian Quindío southwards to northern Peru in a high elevational range between 1,700 and 3,000 m. Although Padrón (2014) does not include C. lycurgus among the studied species, he subsequently managed to obtain good DNA sequences from freshly collected specimens and his results indicate that C. lycurgus would be the sister clade of the Catasticta pieris / eurigania complex and would instead have no relationship with the Catasticta frontina species group (sensu Padrón 2014) in which C. incerta is included (Padrón et al. in prep.).

Remarks on C. lycurgus conservation

The Sierra Nevada de Santa Marta is considered one of the most important centres of endemism and is also well known for the presence of endangered species ( Stattersfield et al. 1998, Alvear et al. 2014), which makes this area of special importance for conservation. This isolated Andean mountain has been one of the national protected areas or National Parks (PNN) of Colombia since 1964, and was declared a UNESCO Biosphere Reserve in 1979 (whc. unesco.org). The Sierra Nevada is one of the Key Biodiversity Areas for Conservation (KBA, keybiodiversityareas. org) and one of the Alliance for Zero Extinction sites (AZE, zeroextinction.org) vital for preventing global species extinctions.

Patterns of highly restricted distribution and endemism have been observed for butterflies ( Takahashi 1976; Shapiro 1979; Adams and Bernard 1977; Adams 1986; Attal and Crosson du Cormier 1996; Torres-Nuñez et al. 1999, Llorente-Bousquets & Le Crom, 2004), and this is especially true for Catasticta . Several butterflies are restricted to this mountain including four species of Catastica that can be found flying along an altitudinal gradient ranging from 1,600 to 3,400m, for example Catasticta rochereaui laurentina , C. chrysolopha adamsi ( Eitschberger & Racheli, 1998) , C. flisa postaurea Brown, 1933 and C. lycurgus .

Notably, several new butterfly taxa have been discovered in the last century in the Santa Marta Mountains (e.g. Adams & Bernard 1977; Steinhauser 2002; Prieto 2011; Llorente-Bousquets & Le Crom 2013, Henao et al. 2015), but no other specimens of this elusive species have been found or at least, are not recorded in accessible collections. Although various researchers and university students have made other studies on butterflies in the Sierra, very few have been published (e.g. Takahashi 1971, 1976; Adams 1973; Montero-Abril & Ortiz-Pérez 2010; Ochoa et al. 2018). The re-discovery of this species means that although scarce and rare, C. lycurgus is not yet extinct as it was feared, as little is known about its biology, its host plant and its conservation status. Its particular habitats in the Sierra Nevada de Santa Marta might be affected by human intervention mainly by agriculture and localised fires and potentially by changes in climate; therefore, a dedicated monitoring programme for the populations of Catasticta lycurgus is urgently needed to protect this species and make use of it as an umbrella species also to protect other endemic and unknown fauna inhabitants of the Sierra Nevada de Santa Marta.

Figures

FIGURE 1. a. Original figure of Euterpe lycurgus from Godman & Salvin (1880). b. Holotype, male specimen dorsal view and labels c. ventral view (B. Huertas, Trustees NHMUK).

FIGURE 2. New male specimens known of C. lycurgus. a. recently collected dorsal view. b. ventral view (M. Bollino) c. female of C. lycurgus illustrated for the first time, dorsal view d. ventral view (C. Prieto) f. male genitalia (M. Bollino).

FIGURE 3. a. Map showing the known collecting localities of C. lycurgus in Colombia. b. forest habitat in San Pedro de la Sierra, road to Las Lagunas at 2,600 m (F. Montero) c-d. live specimens of C. lycurgus.