Dynamine alexae, PEÑALVER & GRIMALDI, 2006

PEÑALVER, ENRIQUE & GRIMALDI, DAVID A., 2006, New Data on Miocene Butterflies in Dominican Amber (Lepidoptera: Riodinidae and Nymphalidae) with the Description of a New Nymphalid, American Museum Novitates 3519 (1), pp. 1-20 : 7-9

Dynamine alexae , new species

Figures 2d , 3 , 4 , 5

MATERIAL: AMNH DR-18-2 (male), holotype, in an amber piece 32 mm length X 22 mm width X 8 mm thickness .

The holotype is incomplete, having lost the anterior part of the body at the surface of the amber (figs. 2d, 3), specifically the head, the anterior part of the thorax, most of the right wings and proximal portions of the left fore- and hind wings. Preserved are only the distal part of the mid- and hind legs, most of the left wings, and abdomen with genitalia. The wings partly preserve the color pattern as tones of brown and are overlapping. The wing surfaces have small to medium gas bubbles trapped during immersion in the resin.

ETYMOLOGY: Patronym in honor of Mrs. Alex Goelet, wife of Mr. Robert G. Goelet, Chairman Emeritus and trustee of the AMNH; for their generous sponsorship of research at the AMNH.

TYPE LOCALITY: Mid-Miocene amber from mines in the Cordillera Septentrional, north and northeast of Santiago, Dominican Republic (Hispaniola).

DIAGNOSIS: According to wing size this new Dynamine species was large. Greatest width of the hind wing larger than greatest width of the fore wing (HW/FW 5 1.19). White median band under both fore and hind wings; costal white subband on fore wing covers M 3 –CuA 1 cell and ends in CuA 1 –CuA 2. Two identical ocelli under hind wing. Continuous, broad, brown postmedial band under hind wing connects the two ocelli, and has a broad, pale brown proximal margin. Two strong, short apical spines at apex of each male valve, one dorsal and one ventral.

DESCRIPTION: Large body size. Forewing (figs. 3a, 4): the preserved portion of forewing indicates a triangular shape (fig. 4), inferred length is 25.80 mm, and the greatest width is 16.11 mm; L/W inferred 5 1.60. White medi- an band on underside broken at level of CuA 1 in two subbands, the apical part of the costal subband covers M 3 –CuA 1 cell and ends in CuA 1 –CuA 2; apical part of anal subband ending in vein CuA 1. Hind wing (figs. 3b, 4): equal in length and width (inferred length is 18.75 mm and greatest width 19.17 mm); L/W inferred 5 0.98. White underside with ocelli and brown bands: underside with paired submarginal ocelli in M 1 –M 2 and CuA 1 – CuA 2, identical and very circular in shape; ocelli connected by broad, brown, continuous postmedial band (this band has a broad pale brown proximal margin); one wide medial brown band preserved has a pale area in the anal region; distal marginal brown band has a longitudinal, pale band for entire length. Proximal part of wing is not preserved. Margin without expansions or tails. Legs: Tibiae with white scales ventrally and two large apical spurs. Mid and hind tarsomeres with four rows of strong setae (two ventral and two ventrolateral), except for tarsomere V, which has two rows of ventral setae (fig. 3d). Length of tarsomeres of midleg: I,? mm; II, 0.73 mm; III, 0.43 mm; IV, 0.29 mm; and V, 0.52 mm. Length of tarsomeres of hind leg: I, 2.29 mm; II, 0.51 mm; III. 0.23 mm; IV, 0.26 mm; and V, 0.46 mm. White scales present on tibiae. Claws unforked. Abdomen dorsally and laterally dark brown and white ventrally. Male genitalia with valves exposed, which are long, flattened, and narrow, and a narrow tubular structure that possibly corresponds to the uncus (figs. 3c, 5); the apex of each valve has two strong, short spines, one dorsal and one ventral.

COMMENTS: The female for this new fossil species is as yet unknown. The reconstruction of the wing color pattern of the preserved areas was partial (fig. 4) because the wings are incomplete, overlapping, and touching.

This specimen belongs to the genus Dynamine due to the combination of the following characters: presence of a white median band on fore- and hind wings, absence of ocelli on the forewing, two ocelli under the hind wing restricted to M 1 –M 2 and CuA 1 – CuA 2 and connected by a broad brown band, hind wing margin without expansions or tails, and a dark brown abdomen that is white ventrally. Some species of Dynamine do not have ocelli, but the two extant species present in the Antilles have ocelli. We include the genus Dynamine in the subfamily Biblidinae and tribe Dynaminini, sensu Oppler and Warren (2003) and Freitas and Brown (2004). The subfamily Biblidinae is one of six groups that have been established by the cladistic analysis of Freitas and Brown (2004).

The family Nymphalidae has been found previously in Dominican amber, represented by a caterpillar placed near the genus Smyrna of the subfamily Nymphalinae ( Hammond and Poinar, 1998) , and thus without any relationship to the new fossil nymphalid specimen.

COMPARISON WITH EXTANT DYNAMINE: The genus Dynamine has very characteristic wing color patterns, especially that of the underside, and for this reason it can be confused only with the genus Lucinia . However, Lucinia has two larger wing ocelli, with each covering more of one cell and not connected by any continuous or broken brown band; and the hind wing margin is not rounded. Röber (1916) superficially segregated the species of Dynamine into three groups: 1) those with little difference in wing pattern between males and females; 2) those with appreciable difference in the patterning of the sexes and without wing ocelli on the underside; and 3) those that differ in the wing pattern of the sexes and which have wing ocelli.

The genus Dynamine is represented in the West Indies by only two species ( D. egaea Fabricius, 1775 and D. mylitta Cramer, 1780 ), both belonging to group 3. Dynamine alexae new species has wing ocelli, but it is unknown whether the sexes were dimorphic and whether the wing uppersides had white spots or bands. Dynamine mylitta has a range from Mexico to Argentina including Cuba, and has a wing pattern very different from that of D. alexae (the former has a white median band under the forewing divided in three small parts and the anterior ocellus on the hind wing is present in CuA 1 –CuA 3). Dynamine egaea has a range from Mexico to Colombia, including Cuba, Hispaniola, and Jamaica, where it purportedly occurs as three subspecies (D.e.calais, D.e.zetes and D.e. egaea , respectively). Dynamine alexae is much larger than all the West Indies forms (25.80 mm estimated forewing length vs. 21– 22 mm; and 18.75 mm greatest hind wing width vs. 12.46 mm; see fig. 4). The broad continuous band that connects the two hind wing ocelli is similar to the band in the extant subspecies D. egaea calais from Cuba and D. e. egaea from Jamaica, and different from the subspecies on Hispaniola that has a narrow, occasionally broken, brown band (fig. 4). In general, the wing color pattern of D. alexae is similar to that of D. egaea on the continent and the West Indies.

The species of Dynamine possessing two ocelli that are not present in the West Indies are: D. ate , D. artemisia , D. onias , D. glauce , D. meridionalis , D. agatha , D. pebana , D. aerata , D. paulina , D. perpetua , D. gisella , and D. zenobia (for wing pattern and other characteristics of these species, see D’Abrera, 1987). Dynamine alexae differs from all of these species in some characters, most obviously in a size significantly larger than that of the first seven species listed. Butterflies in the genus Dynamine are all small-sized (FW length from 13 to 28 mm approx.) and, for this range D. alexae (FW length 5 25.80 mm) equals in size the largest specimens of the largest continental species (e.g., D. hoppi gillotti with a range of FW length from 26 to 28 mm, or D. hecuba from 22 to 24 mm). Furthermore, D. alexae n.sp. has a very broad hind wing compared to the greatest width of its forewing. In extant Dynamine species the greatest width is identical in both wings or even larger in the forewing. In addition, Dynamine aerata has the posterior wing ocellus larger that the anterior one. Dynamine paulina has a very large anterior ocellus that extends over three cells, and the white median band on the underside of the forewing is divided into three small parts (fig. 4). Dynamine perpetua , D. gisella , and D. zenobia have a very small, costal, white subband on the fore wing in comparison with D. alexae .

The male valves in extant Dynamine vary greatly in form. We have compared the valves of D. alexae n.sp. with four species (fig. 5), which differ most in the structure of the apex. The two ‘‘subspecies’’ of D. egaea that were studied from the West Indies have a broad, granulate apex without spines. The continental ‘‘subspecies’’ D. paulina thalassina has three strong, short spines (one dorsal and two ventral) in the apex. Lastly, the continental D. glauce and D. artemisia have two small apical spines.

Figures

Fig. 2. Photomicrographs of Miocene butterflies in Dominican amber. a. Female of Voltinia dramba (AMNH DR-18-1) in ventral view, and complete view of the amber piece. b. Apical tibial spur in the hind leg of the same specimen. c. Dorsal view of the new Voltinia dramba specimen; this butterfly is fossilized with the wings at an angle and so this image was made with six consecutive pictures of the specimen taken at successive focal planes. d. Male of Dynamine alexae n.sp. holotype (AMNH DR-18-2) in dorsal (left) and ventral view (right). Scale bars 5 1 cm in panels a, c, d; 5 0.1 mm in panel b.

Fig. 3. Camera lucida drawings of Dynamine alexae n.sp. (AMNH DR-18-2) in Miocene amber from the Dominican Republic, holotype. a. Body ventral view with the hind-wing color pattern preserved. b. View of the upper side of the forewing, showing the preserved color pattern. c. Male genitalia. d. Hind leg. a–b to same scale.

Fig. 4. Fossil and Recent wing venations and underside color patterns in males of the genus Dynamine. The reconstruction of the underside color pattern of Dynamine alexae n.sp. is adapted from several Recent species of Dynamine (fig. 3 shows the partly preserved color pattern), and the reconstruction of the wing venation (depicted as gray lines) is adapted from the venation of D. egaea zetes from Hispaniola. All specimens are from the AMNH collection. All to same scale.

Fig. 5. Male genitalic valves of fossil and Recent species of Dynamine. All specimens are from the AMNH collection. All to same scale.