Research Article |
Corresponding author: Lucas Campello ( lucas.campellog@gmail.com ) Academic editor: Hume Douglas
© 2022 Lucas Campello, Stephanie Vaz, José R. M. Mermudes, André L. D. Ferreira, Luiz F. L. Silveira.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Campello L, Vaz S, Mermudes JRM, Ferreira ALD, Silveira LFL (2022) Comparative morphology and key to Amydetinae genera, with description of three new firefly species (Coleoptera, Lampyridae). ZooKeys 1114: 131-166. https://doi.org/10.3897/zookeys.1114.77692
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Amydetinae is an exclusively Neotropical subfamily of fireflies, distributed among three genera: Amydetes Illiger, 1807, Magnoculus McDermott, 1964, and Memoan Silveira & Mermudes, 2013. Here, we describe three new species of Amydetinae: two belonging to Amydetes (A. alexi sp. nov. and A. marolae sp. nov.) and one to the previously monotypic Memoan (Me. conani sp. nov.). All three species are known only from the Atlantic Forest in southeastern Brazil. Endoskeletal structures of Memoan and Magnoculus species are described and compared with those of Amydetes for the first time. After studying the type material, Photinus fruhstorferi Pic, 1942 is transferred to Memoan, generating Memoan fruhstorferi comb. nov., and Me. ciceroi Silveira & Mermudes, 2013 syn. nov. is placed as a junior synonym. We also redescribe Magnoculus obscurus Olivier, 1885 and compare it to other species of genus and to other amydetine taxa to identify potential new diagnostic traits for the Amydetinae and its constituting genera. We provide an updated diagnosis for Memoan, illustrations for all four species, and a distribution map for the three new species, as well as a key to adult males of the three amydetine genera, and an updated key to Amydetes species based on males.
Amydetes, Atlantic Forest, Magnoculus, Memoan, Neotropics
Amydetinae (sensu
Amydetes has 21 species across South America (records from Mexico have been disputed:
Both Amydetes and Magnoculus males have flabellate antennae, but the former has between 23 or more antennomeres, while the latter has only 11 (
The lack of zoological studies contrasts with the biotechnological value and potential of the Amydetinae, as some species have been the source of materials for applied research (e.g.,
The amydetines are particularly rich in the Brazilian Atlantic Forest, the second largest tropical forest on the American continent and one of the 25 global biodiversity hotspots (
The Brazilian Atlantic Forest is a global biodiversity hotspot (
The Serra do Mar mountain range, about 1000 km long, constitutes the most outstanding orographic feature of the Atlantic Forest on the South American continent (
The Pedra Branca massif is the largest mountain in the city of Rio de Janeiro. It has a humid, tropical climate without a dry season, with average rainfall of 500–2000 mm from December to March (summer), and 500–1000 mm from June to August (winter). Ilha Grande has a humid, tropical climate with average annual temperatures of 21 °C, ranging between 19.9 °C and 27 °C. Both conservation units are endangered by anthropogenic effects on the Atlantic Forest (
All specimens mentioned here were captured with Malaise traps (190 × 110 cm with 92% ethanol), by active searches in the field, or borrowed from Brazilian natural history museums.
ZMHB Museum für Naturkunde der Humboldt-Universität.
The species distribution maps were made using QGIS v. 2.18.10 (QGIS.org 2017), and the final figures were assembled and edited in Adobe Photoshop CS6.
The material studied consists of 228 specimens from
Photographs were taken using Leica Application Suite CV3 automatic image editing program, and figures edited and assembled using Adobe Photoshop CS6 software.
Family Lampyridae
Memoan
Silveira & Mermudes, 2013; Silveira and Mermudes 2014a: 204;
Vertex straight (Fig.
Male. Head. Capsule 1.5× wider than long (Fig.
Memoan conani sp. nov., mesoscutellum A dorsal; metanotum B dorsal; pterothorax C dorsal; mesoendoesternite D dorsal; metaendoesternite E dorsal. Scale bars: 200 µm (A, D–E); 2 mm (B, C). Note the sutures between mesoventrite/mesanespisternum and mesanepisternum/mesepimeron, the metathoracic discrimen, and irregular flap-like projections on mesendosternum (arrows).
Females and immature stages. Unknown.
The discovery of a second species of Memoan (see below) called for an updated diagnosis for this genus. Memoan is distinguishable from other amydetine genus by their serrate antennae.
Photinus fruhstorferi
Pic, 1942: 16;
Memoan ciceroi Silveira & Mermudes, 2013: 80 syn. nov.
After studying Maurice Pic’s collection at the
Holotype
: Bearing the label: “Espírito Santo. Brasil. ex coll Fruhstorfer.” [aged green label, typewritten]; “TYPE” [aged red label, typewritten]; “R Fruhstorferi” [aged white label, handwritten] (
The specific epithet conani is a masculine noun in the genitive case. The species is named in honor of Mauricio Conan Mendes Correa de Oliveira. Conan was a biology student at the Universidade Federal do Rio de Janeiro, deceased since January 2019, and was a dear friend of the first author.
Labial palp with two palpomeres (Fig.
Male. Coloration. Tegument dark brown (Fig.
Females and immature stages. Unknown.
Twenty-three individuals were collected in hilly areas in the Pedra Branca massif (Fig.
Memoan conani sp. nov. is the second species described in the genus. The differences between species are marked: labial palp (with one palpomere in Me. fruhstorferi comb. nov. and with two palpomeres in Me. conani sp. nov.), which differs morphologically from the all other lampyrids as by
Both species of Memoan occur in remnants of montane forests, separated from one another by a distance of 400 km: Me. fruhstorferi comb. nov. occurs at the Santa Lúcia Biological Station, in Espírito Santo state, whereas Me. conani sp. nov. occurs at the Massif of Pedra Branca, Rio de Janeiro state. Since fireflies are poor dispersers (
Memoan conani sp. nov. were only seen in a narrow spatial and temporal window; that is, males were only observed and collected in a single slope within the limits of the PEPB at an altitude around 400 m a.s.l., and only in April. Likewise, Me. fruhstorferi comb. nov. has only been collected at about 600 m a.s.l., also exclusively in April. Such narrow environmental preferences have been reported for most firefly species occurring in the Atlantic Forest (
Holotype
: Brazil • Rio de Janeiro: Rio de Janeiro: Parque Estadual da Pedra Branca, Núcleo Camorim, Trilha do Açude, 22°58'03.7"S, 43°26'45.7"W; 400 m a.s.l.; ♂; 12 Apr. 2017; L. Silveira, L. Campello, S. Vaz, A.L. Diniz leg. (
The specific epithet marolae is a feminine noun in the genitive case. This species is named in honor of our colleague and friend, Raquel Santos Soares Queiroz, alias “Marola”, who helped us collect the type specimens.
Antennae with scape and pedicel yellowish brown (Fig.
Male. Coloration. Antennae with scape and pedicel yellowish brown (Fig.
Females and immature stages. Unknown.
One hundred forty-six individuals were collected in hilly areas of the Ilha Grande State Park (Fig.
Amydetes marolae sp. nov. is similar to A. bellorum, with which it shares the following combination of traits: labrum connected to fronto-clypeus by membrane; maxillary palpomere IV at least 6× longer than III (up to 7× in A. bellorum, and at least 10× in A. marolae sp. nov.); hypomeron as long as tall; and sternite VIII bisinuate. Nevertheless, Amydetes marolae sp. nov. differs from A. bellorum by: body length (average = 0.67 mm [n = 10, range = 0.6–0.8] in A. marolae and 0.87 mm [n = 10, range = 0.8–1.0] in A. bellorum); pygidium (entirely dark brown in A. bellorum, dark brown with a translucent posterior line in A. marolae sp. nov.).
Amydetes marolae sp. nov. occurs in a mountainous and coastal region of the Serra do Mar range, where it is found between 160 and 660 m a.s.l., with greater abundance in parts below to 400 m a.s.l. Despite extensive sampling (e.g.,
Holotype
: Brazil • Rio de Janeiro: Angra dos Reis: Parque Estadual da Ilha Grande, Pico do Papagaio; 23°09'05.8"S, 44°11'19.9"W; 660 m a.s.l.; ♂; Apr. 2018; L. Campello, L. Silveira, R. Queiroz, S. Vaz leg. (
The specific epithet alexi, is a masculine noun in the genitive case. This species is named in honor of Alex Schomaker Bastos, our dearest friend and biology student at the Universidade Federal do Rio de Janeiro, who was murdered on 8 January 2015.
Antennae with scape and pedicel yellowish brown (Fig.
Male. Coloration. Antennae with scape and pedicel yellowish-brown (Fig.
Females and immature stages. Unknown.
Forty-five individuals were collected in hilly areas in the Pedra Branca State Park (Fig.
Amydetes alexi sp. nov. differs from A. bellorum and A. marolae sp. nov. in the length of maxillary palpomere IV, which is at least 6× longer than III in A. marolae sp. nov. and A. bellorum, but only 3× longer than III in three species: A. itatiaia, A. apicalis, and A. alexi sp. nov. Amydetes alexi sp. nov. differs from A. itatiaia in the length of flabellum of antennomere III (3× longer than pedicel in A. alexi sp. nov., but equal in A. itatiaia). The new species shares the following traits with A. apicalis: pronotum with posterolateral angles pointed but weakly developed; flabellum of antennomere III 3× longer than pedicel; and phallobase asymmetrical. Nevertheless, A. alexi sp. nov. differs from A. apicalis in having antennae with 33–37 antennomeres (37–44 in A. apicalis) and lantern of sternite VI up to 1/2 sternite length (3/5 in A. apicalis). The distribution of A. alexi sp. nov. is apparently restricted to Pedra Branca massif, as it has not been found nearby by our team (e.g.,
Holotype
: Brazil • Rio de Janeiro: Rio de Janeiro: Taquara, Núcleo Pau da Fome, Rio da Fazenda; 285 m a.s.l.; ♂; 6 Aug.–3 Sep. 2017; A. Diniz leg. (
Paratypes
: Brazil • Rio de Janeiro: Rio de Janeiro: Taquara, Núcleo Pau da Fome, Trilha para o açude; 1 ♂; 7–10 May. 2017; L. Silveira, A. Diniz leg. (
1 | Labrum connate to fronto-clypeus, bisinuate, lobes acute; antennal insertions projected ( |
Amydetes bolivari |
– | Labrum free, shape variable, lobes never acute; antennal insertions not projected | 2 |
2 | Elytra with marginal costa with anterior margin rudimentary and not projected ventrally ( |
3 |
– | Elytra with marginal costa with anterior margin developed and projected ventrally ( |
14 |
3 | Antennae at least with 55 antennomeres, maxillary palpomere IV less than 1.5× longer than III | Amydetes solaris |
– | Antennae with 45 antennomeres or less, maxillary palpomere IV at least 2× longer than III | 4 |
4 | Abdominal sternite VIII with posterior margin medially projected | 5 |
– | Abdominal sternite VIII with posterior margin straight or irregular | 9 |
5 | Maxillary palpomere IV at least 6× longer than III | 8 |
– | Maxillary palpomere IV 3× longer than III | 6 |
6 | Lamella I 3× longer than pedicel; pygidium with median lobe rounded | 7 |
– | Lamella I as long as pedicel; pygidium with median lobe beveled | Amydetes itatiaia |
7 | Antennomere III 1/5 longer than pedicel ( |
Amydetes apicalis |
– | Antennomere III 0.5× longer than pedicel (Fig. |
Amydetes alexi sp. nov. |
8 | Pygidium dark brown with posterior 1/3 white, with posterior margin almost straight; phallobase asymmetrical (Fig. |
Amydetes marolae sp. nov. |
– | Pygidium entirely dark brown, with posterior margin bisinuate (i.e., central 1/3 distinctly projected); phallobase symmetrical | Amydetes bellorum |
9 | Pygidium with median lobe at least bisinuate | 10 |
– | Pygidium with median lobe acuminate or rounded (not sinuate) | 13 |
10 | Labrum bisinuate; abdominal sternite VI with lantern almost as long as this sternite | Amydetes vivianii |
– | Labrum straight or slightly concave; abdominal sternite VI with lantern with up to 3/5 length of this sternite | 11 |
11 | Fronto-clypeus with anterior margin almost up to antennal insertions, usually with a discreet median bevel; antennomere III as long as pedicel; pronotum with posterior angle strongly deflexed | Amydetes vagalume |
– | Fronto-clypeus with anterior margin separated from antennal insertions at least 1/2 antennal socket length; antennomere III longer than pedicel; pronotum with posterior angle straight | 12 |
12 | Maxillary palpomere IV 1/3 longer than labial palpomere III; abdominal sternite VI with lantern 0.5× as wide as sternite; abdominal sternite IX 1/3 longer than syntergite, parameres separated; phallus without subapical indentations | Amydetes caetite |
– | Maxillary palpomere IV as long as labial palpomere III; abdominal sternite VI with lantern with less than 1/3 as wide as sternite; abdominal sternite IX 2× longer than syntergite, paramerers touching on the inner margin; phallus with subapical indentations | Amydetes lucernula |
13 | Antennomere III as long as pedicel, pronotum with posterior angle deflexed | Amydetes marajoara |
– | Antennomere III at least 1/5 longer than pedicel, pronotum with posterior angle fairly straight | 15 |
14 | Fronto-clypeus with carinae convergent posterad; pronotum with posterior angle weakly projected; hypomeron 2× longer than tall in lateral view; pygidium with median lobe longer than lateral lobe | Amydetes agnita |
– | Fronto-clypeus without carinae; pronotum with posterior angle well projected; hypomeron 1.5× longer than tall in lateral view; pygidium with median lobe as long as lateral lobe | Amydetes goiana |
15 | Antennomere III with flabellum as long as antennomere III | 16 |
– | Antennomere III with flabellum notably longer antennomere III | 17 |
16 | Pronotum with posterior angle deflexed; maxillary palpomere VI 4× longer than III; indentation between lobes median and lateral rudimentary | Amydetes discissa |
– | Pronotum with posterior angle straight; maxillary palpomere VI 2× longer than III; pygidium with lateral lobe weakly developed, acute, indentation between lobes median and lateral weakly developed | Amydetes manezinha |
17 | Abdominal sternite V with posterior margin concave ( |
18 |
– | Abdominal sternite V with posterior margin straight ( |
19 |
18 | Gular sutures separated by 1/2 maxillary palpomere IV width; hypomeron 2× longer than tall in lateral view; abdominal sternites with posterior margin strongly concave; abdominal sternite VI with lantern as long as 3/5 sternite length and as wide as 3/5 sternite width | Amydetes fastigiata |
– | Gular sutures separated by maxillary palpomere IV width; hypomeron 1.5× longer than tall in lateral view; abdominal sternites with posterior margin concave; abdominal sternite VI with lantern almost as long as wide as sternite | Amydetes fucata |
19 | Pronotum with posterior angle obtuse | Amydetes lucioloides |
– | Pronotum with posterior angle acute | 20 |
20 | Maxillary palpomere IV 2× longer than III | Amydetes plaumanni |
– | Maxillary palpomere IV 4× longer than III | Amydetes luzecu |
(nec Megalophthalmus Leach, 1830 [Crustacea])
Megalophthalmus obscurus Olivier, 1885: 146.
Magnoculus obscurus
(Olivier, 1885)—
Elytra with inner margin dehiscent (i.e., sinuose inner margin) at posterior 1/3 (Fig.
Magnoculus obscurus, mesoscutellum A dorsal; metanotum B dorsal; pterothorax C dorsal; mesoendoesternite D dorsal; metaendoesternite E dorsal. Scale bars: 500 µm (A, D–E); 1 mm (B, C). Note the sutures between mesoventrite/mesanespisternum and mesanepisternum/mesepimeron; the metathoracic discrimen; and flap-like projections on mesendosternum (arrows).
Male. Coloration. Antennae dark brown (Fig.
Females and immature stages. Unknown.
Fourteen individuals were collected in hilly areas in Ilha Grande State Park between 160–345 m a.s.l. Six specimens were collected using Malaise traps: three at 160 m a.s.l. in August 2017, December 2017, and January 2018; two at 170 m a.s.l. in November 2017 and April 2018; one at 345 m a.s.l. in August 2017. In addition, nine specimens were collected by active searches in July 2017 (five specimens), September 2017 (two specimens), and July 2018 (two specimens). Ma. obscurus has a yellowish-green glow, and males often fly between 0.1 and 4 m a.s.l., sometimes reaching up to approximately 7 m a.s.l. towards the forest canopy. Adults are apparently active in the early twilight hours. About five to ten males were observed flying close together in the same visual field.
Magnoculus obscurus is the second species of genus to be redescribed (see Zaragoza-Caballero 1995 for redescription of Ma. cf. guatemalae). The few species recently studied and illustrated in detail by
A comparison of Ma. obscurus with species described and illustrated in detail by
Bearing the labels: “SYNTYPE. Megalophthalmus. obscurus. Olivier, 1885.
Olivier mentioned examining specimens in his and also in Oberthür’s collections. It is unclear how many specimens Olivier examined, and we did not have the opportunity to carefully look for them at the
Brazil • Rio de Janeiro: Angra dos Reis: Parque Estadual da Ilha Grande), Pico do Papagaio; 4 ♂; 29 Jul. 2017; L. Silveira leg. (
1 | Antenna with at least 23 antennomeres (Figs |
Amydetes |
– | Antenna with 10 or 11 antennomeres (Figs |
2 |
2 | Antenna flabellate (Fig. |
Magnoculus |
– | Antenna serrate (Fig. |
Memoan |
Since
When comparing the three amydetine genera, a few interesting similarities and differences are evident (summarized in Table
Amydetes | Memoan | Magnoculus | |
---|---|---|---|
Antennae | Flabellate, with 23–62 antennomeres; antennal sockets reniform | Serrate, with 10 antennomeres; antennal sockets with inner margin straigth and outer margin rounded | Flabellate, with 11 antennomeres; antennal sockets rounded |
Fronto-clypeal suture | Connected by membrane | Connate | Connate |
Mouthparts | Labial palp with three palpomeres | Labial palp with one or two palpomeres | Labial palp with three pelpomeres |
Gula | Gular margins separated by length of labial palpomere II | Gular margins contiguous | Gular sutures contiguous |
Pronotum | semilunar, without two sclerotized raised tubercles on the 1/2 posterior; disc with fine punctures | Rectangular; two sclerotized raised tubercles on the 1/2 posterior; disc with deep punctures | Semilunar; two sclerotized raised tubercles on the 1/2 posterior, with or without deep punctures |
Elytron | Outer margin emarginate | Outer margin rounded | Outer margin rounded; with three raised costae |
Thorax | Metathoracic discrimen as long as 3/4 of sternum length | Metathoracic discrimen as long as 1/2 of sternum length | Metathoracic discrimen as long as 3/4 of sternum length |
Aedeagus | Phalobase symmetrical or asymmetrical | Phalobase symmetric | Phalobase symmetric or asymmetric |
The pronotum has two sclerotized raised tubercles in Magnoculus and Memoan, which are absent in Amydetes, and there are wide punctures with little interstices in the three genera, although in Amydetes these are restricted to the pronotal expansions. The pronotum is semilunar in Magnoculus and Amydetes, but rectangular in Memoan. The eyes are ventrally close-set in both Memoan spp., but only in some species of Amydetes and Magnoculus. Finally, the phallobase is symmetrical in both Memoan spp., but may be symmetrical or asymmetrical in Amydetes and Magnoculus species.
Memoan conani sp. nov., Amydetes marolae sp. nov., and Amydetes alexi sp. nov. are three new additions to the subfamily Amydetinae recently collected in protected areas. All three species appear to have narrow spatial and temporal distributions and are thought to be endemic to their localities, as known for other amydetine species (e.g.,
Authorization to carry out research in the state parks mentioned in this work was granted by INEA 056/2016; E-07/002.11741/2016 and Chico Mendes Institute for Biodiversity Conservation, Ministry of the Environment, Brazil, under Scientific Collection/Research License 21409-5. We thank the staff of the aforementioned conservation units for all the support given to field research and members of the Entomology Laboratory and Insect Ecology Laboratory, UFRJ. We would like to thank everyone who effectively helped to carry out this work, especially A. Katz for all the help with collection, observation and photographing, W. Grande for all the support and assistance in the field, and the four reviewers who contributed much with their reflections and comments on the manuscript: Viridiana V. Badillo, André S. Roza, Cisteil X. Pérez-Hernández, and Hume Douglas. We also thank
Figures S1, S2
Data type: pdf file
Explanation note: Original labels of the type species of Magnoculus obscurus and Memoan fruhstorferi comb. nov. deposited in the