Corresponding author: Philipp Hoenle (
Academic editor: Brian Lee Fisher
One of the largest species in its genus,
Describimos una especie nueva, entre las más grandes conocidas del género
Hoenle PO, Lattke JE, Donoso DA, von Beeren C, Heethoff M, Schmelzle S, Argoti A, Camacho L, Ströbel B, Blüthgen N (2020)
Members of the genus
Ecuador, with at least 18
We describe a surprisingly large new species, i.e.,
Specimens of
The worker holotype, two worker paratypes, and one paratype queen were mounted with water-soluble insect glue on the tip of an insect needle and imaged using the Darmstadt Insect Scanner (
Additional stacking pictures were taken with a Canon EOS 7D with a MPE 65mm lens (Canon, Tokyo, Japan) and a Keyence VHX-5000 (Keyence Deutschland BmH, Neu-Isenburg, Germany) with a Z20 lens. Stacking pictures were assembled with Helicon Focus Version 7 (Helicon Soft Ltd., Kharkiv, Ukraine) software, and further edited with Adobe Photoshop CS6 13.0 (Adobe Inc., San Kaso, CA, USA).
Morphological measurements of three workers (the holotype and two paratypes) and one alate queen were performed using 3D-models embedded in PDF files obtained with
All measurements are given in millimeter.
The collection abbreviation is taken from
We sequenced the classical mitochondrial barcode region for animals, a 658-base pair (
To evaluate whether
Published and newly acquired data from this study were then analyzed together. We first performed several quality checks. Sequences lacking species identifications, sequences containing ambiguous bases, and sequences smaller than 400bp were sorted out. We then used the MUSCLE algorithm (
We used a Neighbor-Joining (
This key is a modification of the keys of
From the
For easy identification, we recommend to point mount workers of
1 | Petiole pedunculate to subpedunculate, in lateral view the anterodorsal margin of the gaster forms a single convexity that ascends posterad at approximately 45° (Fig. |
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– | Petiole sessile, not subpendunculate, in lateral view the anterodorsal margin of the first gaster segment forming a much steeper slope (>45°) with a more or less distinct vertical anterior face (Fig. |
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2 | Dorsal surface of the head with deep striation that reaches to the nuchal collar; color uniform ferruginous to dark red |
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– | Posterior third to half of dorsal surface of head smooth and shining; color variable |
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3 | Without metasternal process; mesonotum almost hairless; larger body size ( |
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– | With metasternal process; dorsal surface of mesonotum covered in small erect setae; smaller body size ( |
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4 | Dorsal surface of head distinctly striate to or nearly to the nuchal carina |
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– | Posterior third to half of dorsal surface of head smooth and shining, or nearly so |
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5 | Disc (dorsal surface) of first gastric segment predominantly smooth, punctulate, alutaceous, or reticulate; striation absent, or if present, mixed with other sculpture and distinct only on the posterior half of the disc |
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– | Disc of first gastric segment distinctly and evenly striate over its entire surface, at least as seen from dorsal view |
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6 | Mesonotum longitudinally striate |
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– | Mesonotum prevailingly transversely striate |
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7 | Head more or less bright red (frontal area often infuscate), contrasting with blackish-brown body and yellow legs; size medium |
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– | Color combination otherwise; if head is distinctly red, then trunk is red also, or legs are dark |
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8 | Sternum immediately in front of and between metathoracic coxae produced as a slender, acute pair of teeth or spines; disc of first gastric segment densely and finely shagreened and pubescent, usually opaque; body brown, legs yellow to brown |
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– | Sternum in front of metathoracic coxae with a low transverse ridge, sometimes notched in the middle or bilobed, but not produced as acute, paired teeth |
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9 | Petiole predominantly smooth and shiny, with the anterior border erected or slightly convex, and the apex produced into a large spine with posterior orientation; head, mesosoma and petiole with a clear red coloration and the gaster dark brown |
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– | Petiole differently shapes, color combination varies |
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10 | Anterior face of petiolar node as seen from the side rising steeply from anterior margin, then passing through an obtuse angle into a long section concave in outline to the root of the apical spine; labial palpi 4-merous |
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– | Petiole differently shaped |
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11 | Metasternal process like an arc with or without middle division; petiole smooth or a little striate, with both the anterior and posterior margin convex; the petiolar spine forms gradually, without clear distinction from petiole | |
– | Metasternal process absent, bilobed or triangular; petiole differently shapes or if both sides convex, with a clear differentiation of the spine from the rest of the petiole |
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12 | Metasternal process absent |
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– | Metasternal process bilobed or triangular |
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13 | Black coloration; Node of petiole with a pair of prominent posterolateral tumosities at about mid-height and without striation; apex as seen from side abruptly narrowed to an axially erect, acute tooth |
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– | Node of petiole without paired posterolateral tumosities; Posterior face of petiole less concave, with short petiolar spine (0.1 mm) |
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14 | Anterior margin of petiole at least weakly convex |
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– | Anterior margin of petiole basal of the node concave or straight |
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15 | Metasternal process completely bilobed; color generally dark |
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– | Metasternal process formed by an obtuse wide lobe followed by a transverse flange which is produced into a triangular process; usually light color |
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16 | Petiole strongly transversely striate, with a clearly differentiated spine; small species (TL 8.6–9.35 mm) |
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– | Petiole without or only weak striation and with spine not clearly differentiated from the petiole; large species (TL 12 mm) |
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17 | First gaster segment with only one type of sculpture, which is either punctulate or striate |
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– | First gaster segment with a combination of punctulate and reticulate sculpture |
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18 | First gaster segment punctulate on its entire surface |
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– | First gaster segment striate on most of its surface, at least in dorsal view |
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19 | Transverse striation patterns on the gaster; large and slender |
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– | Longitudinal striation on the dorsal gaster surface |
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20 | Mesonotum strongly convex, but broadly sulcate and longitudinally striate on at least the anterior half near midline |
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– | Mesonotum gently but evenly convex, transversely striate |
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21 | Mesonotum with longitudinal striation |
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– | Mesonotum with transverse striation |
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22 | Ocular prominences each produced anterolaterally into a stout, acute, oblique, toothlike process |
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– | Ocular prominences bluntly rounded, as usual |
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23 | Antennal scapes very short, not reaching posterior border of head in full-face view; very small species with broad head |
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– | Antennal scapes surpassing posterior border of head viewed full-face |
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24 | Apex of mandible with only 2 large teeth (intercalary tooth lacking) |
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– | Apex of mandible with 3 teeth |
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25 | Mesepisternum with a prominent, narrowly rounded anteroventral lobe projecting conspicuously on each side when trunk is viewed from above |
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– | Mesepisternum with at most a low, inconspicuous convexity on its anteroventral margin |
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26 | Petiole clearly differentiated spine; larger species ( |
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– | Both faces of petiole converge into a thick spine that is flattened laterally; smaller species ( |
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1 | Pecíolo en vista lateral pedunculado a semi-pedunculado, el perfil anterodorsal del primer segmento del gáster forma una convexidad contínua con una pendiente de aproximadamente 45° (Fig. |
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– | Pecíolo sésil, no semi-pedunculado, el perfil anterior del primer segmento del gáster relativamente vertical y bien diferenciado del perfil dorsal, con una pendiente mayor de 45° (Fig. |
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2 | Superficie dorsal de la cabeza con estrías que llegan hasta la carena nucal, color ferruginoso |
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– | Superficie dorsal de la cabeza con estrías que tan sólo ocupan de la mitad a dos tercios de la región anterior de la cabeza, color es variable |
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3 | Espacio entre las coxas posteriores liso, sin proceso ni estrías; mesonotum casi sin pelos; gran tamaño (LC > 4 mm). Hormiga del suelo |
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– | Espacio entre las coxas posteriores con un proceso bilobulado y siempre estriado; mesonotum con muchos pelos; más pequeña (LC < 3.8 mm). Hormiga arbórea |
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4 | Superficie dorsal de la cabeza con estrías que llegan hasta la carena nucal o muy cerca de ésta |
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– | Superficie dorsal de la cabeza con estrías que tan sólo ocupan de la mitad a dos tercios de la región anterior de la cabeza |
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5 | Primer segmento del gaster predominantemente liso y brillante, opaco o suavemente reticulado |
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– | Primer segmento del gaster con escultura que puede ser de un sólo tipo o una mezcla de varios (estríado, punteado, estríado- punteado) |
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6 | Mesonoto estríado longitudinalmente |
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– | Mesonoto estríado transversalmente |
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7 | Cabeza de color rojo claro que contrasta con cuerpo marrón oscuro a negro y extremidades amarillas |
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– | Diferente combinación de color; si la cabeza es rojo claro, entonces el mesosoma debe ser también rojo o las extremidades de un color oscuro; o la cara anterior del pecíolo es recta o cóncava |
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8 | Metaesterno, exactamente entre las coxas posteriores posee un par de espinas o dientes agudos; primer segmento del gaster reticulado, usualmente opaco; cuerpo marrón, extremidades de color amarillo a marrón |
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– | Metasterno sin o con proceso, el cual puede ser bilobulado, dividido en la mitad o redondeado |
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9 | Pecíolo predominantemente suave y brillante, con borde anterior recto o ligeramente convexo, el ápice de éste se estrecha formando una espina larga, delgada que está dirigida posteriormente; cabeza, mesosoma y pecíolo de color rojo claro y gaster marrón oscuro |
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– | Pecíolo de diferente forma; combinación de color variada |
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10 | Cara anterior del nodo peciolar se levanta casi verticalmente desde el margen anterior, luego pasa por un ángulo obtuso a una sección larga y cóncava que forma una espina apical |
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– | Pecíolo con forma diferente |
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11 | Proceso metasternal como un arco con o sin división en el medio; pecíolo suavemente o poco estríado, la cara anterior es convexa, al igual que la posterior; la espina del pecíolo se va formando gradualmente, lo cual hace que no sea claramente diferenciada de éste | |
– | Proceso metasternal ausente, bilobulado o triangular; pecíolo con diferente forma o si ambos lados son convexos hay una espina claramente diferenciada del resto del pecíolo |
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12 | Proceso metasternal ausente |
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– | Proceso metasternal bilobulado o triangular |
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13 | Color negro; nodo del pecíolo con un par de prominencias posterolaterales y sin estrías; ápice en vista lateral se estrecha hasta formar un diente agudo axialmente erguido |
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– | Color claro; nodo del pecíolo sin prominencias o si las posee tiene estrías; cara posterior del pecíolo al menos débilmente cóncava, la espina peciolar es corta (0.1mm) |
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14 | Cara anterior del pecíolo al menos débilmente convexa |
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– | Parte anterior basal del nodo peciolar cóncava o recta |
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15 | Proceso metasternal completamente bilobado; color oscuro generalmente |
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– | Proceso metasternal formado por un lóbulo ancho obtuso, seguido de un reborde transverso que se observa como un proceso de forma triangular; color claro generalmente |
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16 | Pecíolo fuertemente estríado transver- salmente, presenta una espina claramente diferenciada, especies pequeñas (8.6- 9.35mm) |
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– | Pecíolo sin estrías o suavemente estríado, la espina no está claramente diferenciada del pecíolo, especies grandes (TL 12 mm) |
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17 | Primer segmento del gaster con un sólo tipo de escultura, que puede ser punteada o estriada |
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– | Primer segmento del gaster con una combinación de escultura punteada y reticulada |
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18 | Primer segmento del gaster punteado a lo largo de toda su superficie |
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– | Primer segmento del gaster estríado a lo largo de toda su superficie, al menos en vista dorsal |
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19 | Estrías transversales curvas en el gaster; especies grandes y delgadas |
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– | Estrías longitudinales en el dorso del gaster |
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20 | Mesonoto fuertemente convexo, pero fuertemente surcado y estríado longitudinalmente al menos en la mitad del área de la parte |
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– | Mesonoto suave pero uniformemente convexo, estríado transversalmente |
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21 | Mesonoto estríado longitudinalmente |
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– | Mesonoto estríado transversalmente |
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22 | Prominencias oculares con un proceso agudo, oblicuo a manera de diente |
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– | Prominencias oculares redondeadas |
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23 | Escapos antenales muy cortos que no alcanzan el borde posterior de la cabeza |
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– | Escapos antenales sobrepasan el borde posterior de la cabeza |
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24 | Dos dientes grandes en el ápice de la mandíbula |
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– | Tres dientes en el ápice de las mandíbulas |
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25 | Mesepisterno con un lóbulo anteroventral redondeado y prominente que se proyecta a los lados del mesosoma en vista dorsal |
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– | Mesopleura con una convexidad inconspicua en su margen anteroventral |
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26 | Pecíolo convexo en ambas caras, con una espina delgada claramente diferenciada; más grande (LC>2.8 mm) |
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– | Pecíolo con ambas caras convexas, las cuales convergen en una espina gruesa y aplanada lateralmente; más pequena (LC<2.8 mm) |
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Lateral view of petiole and first gaster segment of
Complete list of localities in Suppl. material
Ecuador • Esmeraldas, Reserva Río Canandé;
3D models of
Ecuador • 1 worker; Esmeraldas, Reserva Río Canandé;
Ecuador • 3 workers; Esmeraldas, Reserva Rio Canande;
Ecuador • 2 workers; Esmeraldas, Reserva Río Canandé;
Ecuador • 4 workers; Esmeraldas, Reserva Río Canandé;
Ecuador • 4 workers; Esmeraldas, Reserva Río Canandé;
Ecuador • 2 queens; Esmeraldas, Reserva Río Canandé;
Ecuador • 1 queen; Esmeraldas, Reserva Río Canandé;
Ecuador • 1 queen; Esmeraldas, Reserva Río Canandé;
Ecuador • 1 queen; Esmeraldas, Kumanii Lodge near Cotocachi-Cayapas Reserve;
1 holotype worker (PE39_01), 1 paratype worker (PE23_01), 1 paratype worker (PE36_01), 1 paratype queen (PE24_01)
Paratype workers
Long (TL > 17 mm), but slender, ferruginous to yellow brown body with striae on cephalic dorsum from antennal insertions to vertex, mandible with over 15 pre-apical teeth and denticles, pronotal dorsum with concentric to transverse striae. Petiole strongly pedunculate with posteriorly inclined apical spine, gaster smooth and shining.
Head elongate in dorsal view, anterior and posterior margins approximately of same width, posterior cephalic margin mostly transverse; head widest across eyes, at anterior one-third of head length; lateral cephalic margin posterior to eye sinuous. Median furrow deep, extends anterad to antennal fossa where it fades; occipital ridge distinctly delineated by antennal fossa, extending posteromedially, joining broad ridge that runs parallel to median furrow. Extraocular furrow broad and shallow, temporal prominence broad and weakly elevated. Cephalic surface with well-defined striae that diverge posterad from between frontal carina, reaching vertex, striae fade away on most of lateral cephalic surface with some striae reaching posteroventral cephalic surface. Ocular ridge smooth closest to eye and striate towards cephalic median region. Cephalic dorsal surface anterad of eye and between eye and antennal sclerite mostly smooth. Scape slender and slightly arched,
Median clypeus mostly smooth and shining, posteriorly projecting as flattened triangular surface between frontal carina; carina defines narrow elevated region that descends posteriorly and extends to antennal fossa; frontal carina narrow, width not greater than scape width; carina steeply elevated over posteromedian clypeal surface. Ventral cephalic surface mostly smooth and shining. Labium drop-shaped, anteroventral surface very convex, PF 4,4. Buccal cavity with lateral hypostomal tooth. Mandibular masticatory margin with basal row of six denticles and eleven blunt triangular, relatively short teeth apicad of denticles. One or more teeth closest to apex may be broken. Mandibular apex tridentate, ventral tooth with basal tooth. Mandibular dorsal surface mostly smooth, with sparse piligerous punctulae, but dorsolaterally with abundant punctulae, ventral surface smooth and shining.
Pronotal dorsum with concentric striation that become progressively transverse and elongate medially towards posterior margin, in lateral view striae appear anteriorly transverse, medially curving and U-shaped, posteriorly oblique to almost vertical. Posterolateral pronotal margin with short convex lobe. Mesosoma relatively slender and elongate, in lateral view pronotal dorsal margin straight to weakly convex, forming a posteriorly ascending slope, mesonotum anterior margin slightly higher than posterior pronotal margin, mesonotal dorsal margin mostly straight to weakly convex, descending to metanotal groove. Dorsal mesosomal margin between metanotal sulcus and metanotal spiracle forms brief convexity, propodeal anterodorsal margin brief and convex, dorsal margin mostly straight, three times longer than declivity, declivity forms blunt obtuse angle with dorsal margin. Propleuron mostly smooth and shining with narrow transverse band of sparse weak rugulae anteriorly and posteriorly.
Mesonotum with transverse striae that extend uninterrupted laterally to anepisternum and ventrally to mesosternum, katepisternum mostly smooth and shining except for sparse striae anteriorly and posteroventrally. Bulla of metathoracic spiracle semispherical, weakly sculpted, opening shaped as transverse slit. Propodeum and metanotum transversely striate. Mesometapleural suture distinct, propodeal- metapleural suture weakly impressed. No carina or visible suture between mesopleuron and mesosternum, mesosoma in hypothetical cross-section at mid-length forms relatively uniform ovoid. Mesosternum with median longitudinal region raised as low and broad convex ridge; metasternal process bidentate, teeth short and blunt. Propodeal spiracle slit-shaped, transverse to oblique, not elevated.
Petiole in lateral view slightly pedunculate, node shaped as posteriorly sloping cone with acute apical needle, anterior node margin weakly convex, posterior margin vertical, straight to weakly convex; anteroventral process prominent, triangular; node smooth and shining. Abdominal tergite 3 in lateral view with anterodorsal margin forming single convexity to posterior margin, ascending posterad at approximately 45°; ventral margin of tergite 3 briefly concave at prora, then broadly convex and mostly at the same level as prora. Constriction between abdominal segments 3, 4 weak to negligible; gaster smooth and shining.
Coxae mostly smooth with abundant minute piligerous punctulae, punctulae denser on tibiae. Protibial apex with single seta, spur with basal translucent lamella. Probasitarsus with row of short, stiff hairs and parallel row of short setae opposite spur. Meso and metatibial apex each with two spurs, one pectinate, one simple; each also with 3 setae, each seta widely separated from each other. Body pilosity generally short and scattered with little pubescence; dorsal surface with few standing hairs: one on head where antennal fossa and nuchal carina almost meet, few on gastral sterna. Head and mandibular dorsum with sparse appressed pubescence, hairs straight on mandible and arched on head. Mandibular ventral surface next to masticatory margin with row of five flagellate long hairs plus two long trigger hairs at base. Scape with dense appressed pubescence, no standing hairs. Mesosoma with sparse appressed to subdecumbent small hairs, node with longer hairs; gaster mostly with sparse short, appressed to decumbent hairs with suberect hairs towards posterior end of gaster. Mandible and other buccal appendages, antenna, tibiae, and tarsi ferruginous brown to brown. Body mostly ferruginous to brownish yellow, head dark anterad and gaster darker posterad; trochanters and apex of femora tend to be darker.
Mesosoma developed for wings, head with three ocelli. Queen with larger dimensions than worker:
Compared to
While the body size of
When using the key to Neotropical
We successfully amplified DNA barcodes of two
Neighbor-Joining (
Workers of
On the 11th of February 2019 we collected what looked like a complete nest in a fallen branch (Fig.
Nest of
Colony of
We kept the colony for three months (11 February–15 April 2019) for further observations. The colony accepted various smaller insects as food, including flies, crickets, and termites. However, insects larger than 2 cm (e.g., large cicadas, moths, large crickets) were usually not accepted. Furthermore, the colony had
We here formally described the species
Sequence similarities of
Our species description is accompanied by 3D scans of three workers and a queen (Fig.
The new species was discovered in the Chocó-Darién bioregion in Ecuador, probably one of the most biodiverse regions on earth, and at the same time one of the most threatened ones (
We thank the Fundacíon Jocotoco and the associated Tesoro Escondido for logistic support and their permission to do research on their forest properties and in particular Martin Schaefer for initiating this collaboration. We like to thank especially the local support from the park staff in the Canandé and Tesoro Escondido reserve, that made the field collection easier and made two great field stays possible: Bryan Amayo, Alcides Zombrano, Roberto de la Cruz, Jorge Zambrano, Amado de la Cruz, Yadria Giler, Patricio Encarnación, and Vanessa Moreira.
The species name was auctioned for a good cause at the anniversary of the Rainforest Trust, and we like to thank them and the highest bidder for this opportunity to aid in the conservation efforts and research in the Chocó area.
We would like to thank Brian L. Fisher as well as three reviewers for helpful comments on the manuscript. We acknowledge support by the German Research Foundation and the Open Access Publishing Fund of Technische Universität Darmstadt. PH was supported by a scholarship from the German National Academic Foundation. CvB received funding from the German Research Foundation (DFG: BE 5177/4-1 & BE 5177/4-2). SS was funded by the German Federal Ministry of Education and Research (BMBF), project NOVA (05K2016 / 05K16RDD).
Video of
multimedia
Rendered video of the 3D scan of the holotype specimen.
Video of
multimedia
Rendered video of the 3D scan of the queen paratype.
Table with measurements
measurements
Table containing all available measurements of holo- and paratypes of
DNA barcode information
barcode information
Table containing a list of the barcoded individuals and their associated collection and repository data.
Used BOLD barcodes
BOLD accession numbers
List of BOLD barcode accession numbers used in this study.
newick tree file
Newick file of the
Additional detail images of
multimedia
Contains additional detail pictures of the new species.
Additional 3D scans of two more paratype workers
multimedia
This file contains two additional 3D scans of two more paratype workers.
Table of localities
occurence
Table of known