Research Article |
Corresponding author: Kelly B. Miller ( kbmiller@unm.edu ) Academic editor: Mariano Michat
© 2020 Kelly B. Miller.
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:
Miller KB (2020) Two new species of Agaporomorphus Guignot from Suriname (Coleoptera, Adephaga, Dytiscidae, Copelatinae). ZooKeys 923: 51-63. https://doi.org/10.3897/zookeys.923.48337
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Two new species are described in the Neotropical genus Agaporomorphus Guignot from Suriname: A. hamatocoles sp. nov. and A. tortus sp. nov. The species are included in a phylogenetic parsimony analysis of 13 morphological characters and all 12 known species. Two equally parsimonious arrangements are found with the only difference a rearrangement among the A. knischi clade. Agaporomorphus tortus belongs to the A. dolichodactylus group based on presence of an elongate, club-like lobe on the dorsal, basal surface of the male median lobe and long, subsinuate male mesotarsal claws and a small lobe at the apex of male mesotarsomere V. Agaporomorphus hamatocoles does not belong to a known species group and is phylogenetically isolated lacking synapomorphies characterizing the other groups, so the species is placed in its own species group. Male genitalia are illustrated for the new species and redrawn for all the species of the A. dolichodactylus group, and male mesotarsal claws are illustrated for A. tortus and redrawn for other members of the A. dolichodactylus group. New distribution records are reported for Suriname for the species A. colberti Miller and Wheeler and A. pereirai Guignot.
Se describen dos nuevas especies en el género neotrópico Agaporomorphus Guignot de Surinam: A. hamatocoles sp. nov. y A. tortus sp. nov. Las especies se incluyen en un análisis de parsimonia filogenética de 13 carácteres morfológicos y las 12 especies conocidas. Se encuentran dos arreglos igualmente parsimoniosos, con la única diferencia de un reordenamiento entre el clado de A. knischi. Agaporomorphus tortus pertenece al grupo A. dolichodactylus basado en la presencia en el macho de un lóbulo alargado, que parece un palo en la superficie dorsal, al base del lóbulo mediano ; las largas garras mesotarsales subsinuosas; y un lóbulo pequeño en el ápice del mesotarsómero V. Agaporomorphus hamatocoles no pertenece a ningún grupo de especies conocidas, está aislada filogenéticamente y carece de sinapormorfias que caracterizan los otros grupos, así que la especie se coloca en su propio grupo . Los genitales de los machos se ilustran para las nuevas especies y también para todas las especies del grupo A. dolichodactylus. Las garras mesotarsales de los machosse ilustran para A. tortus y también para los otros miembros del grupo A. dolichodactylus. Se informan nuevos registros de distribución para Surinam para las especies A. colberti Miller y Wheeler y A. pereirai Guignot.
Agaporomorphus, diving beetle, South America, taxonomy, water beetle
New species of Agaporomorphus Guingot have been discovered regularly as collecting has continued in new areas of South America (
Species of Agaporomorphus are known only from lowland tropical South America (
The new species are based on specimens from the Snow Entomological Collection, University of Kansas, Lawrence, Kansas, USA (
Measurements were acquired using an ocular scale on a Zeiss Discovery V8 dissecting microscope at 50× magnification. Measurements include:
TL total length;
GW greatest width across elytra;
PW greatest pronotal width;
HW greatest width of the head;
EW distance between the eyes;
FL greatest length of the metafemur;
FW greatest width of the metafemur.
The ratios TL/GW, HW/EW, and FL/FW were also calculated to provide an indication of overall shape, eye size, and leg segment size.
The new species were coded for the 12 characters described by
Character 13. Apical lobe on male lateral lobe; (0) not extremely long and slender (Fig.
Data matrix of assigned states of characters for 12 species of Agaporomorphus and generalized outgroup based on numerous examined taxa (e.g., Copelatus Erichson, Madaglymbus Shaverdo and Balke, Lacconectus Motschulsky, and Exocelina Broun species). Character 01 coded as additive (others binary). Characters match numbered characters from
Species | 0000000001111 |
---|---|
1234567890123 | |
Outgroup | 0000000000000 |
A. hamatocoles | 0000000000000 |
A. pereirai | 0000010000000 |
A. knischi | 0000011111110 |
A. tambopatensis | 0000001111010 |
A. colberti | 0000001111110 |
A. julianeae | 0000001101110 |
A. silvaticus | 0000001000010 |
A. sharynae | 0000011000010 |
A. grandisinuatus | 1010000000001 |
A. mecolobus | 2111100000001 |
A. dolichodactylus | 2111100000001 |
A. tortus | 2111100000001 |
Suriname, Sipaliwini District, Sipaliwini Savannah Nature Reserve, Four Brothers Mountains, 2.005700N, 55.969151W, 337 m.
This species does not share many features with other members of the genus and does not have modified antennomeres, modified male mesotarsal claws or a lobe on the apex of mesotarsomere V, it lacks a stridulatory apparatus on the abdomen and metaleg, and lacks a triangular process at the apical margin of visible sternite V of the abdomen. Unique features of A. hamatocoles are the strongly hooked male median lobe (Fig.
Measurements (N = 3). TL = 2.8–3.2 mm, GW = 1.4–1.6 mm, PW = 1.2–1.3 mm, HW = 0.8 mm, EW = 0.5–0.6 mm, FL = 0.7–0.8 mm, FW = 0.2–0.3 mm, TL/GW = 1.9–2.0, HW/EW = 1.5–1.6, FL/FW = 2.9–3.4. Body shape elongate oval, evenly and shallowly curved along lateral margins, curvature continuous between pronotum and elytron.
Coloration. Head and pronotum dark orange. Elytron dark orange throughout except transverse basal band light orange. Ventral surface orange, similar in coloration throughout but legs distinctly lighter in color.
Sculpture and structure. Head shiny, very finely microreticulate comprised of small isodiametric cells; eyes small (HW/EW = 1.5–1.6). Pronotum shiny, similar microreticulation to head; lateral margin slightly curved, extremely finely beaded, bead absent at anterior angle. Elytron with lateral margin shallowly curved; surface shiny, microreticulation extremely fine, apical half with numerous extremely fine punctures. Prosternum elongate, carinate, prosternal process short, strongly carinate medially. Metaventer and metaventral wings smooth and shiny, with very dense, extremely fine microreticulation. Metacoxa smooth and shiny, similar in microsculpture to metaventer; metacoxal lines distinct, region between metacoxal lines narrow medially; metafemur not unusually broadened (FL/FW = 2.9–3.4).
Male genitalia.
Median lobe exceptionally complex in shape, strongly asymmetrical; in lateral aspect broad basally, irregularly shaped, apically narrowed with apex dramatically hooked, curved anteriorly on dorsal surface with elongate apex directed posteriorly, curved portion elongate, slender and apically narrowly rounded (Fig.
Agaporomorphus species, male genitalia. 1–3 A. hamatocoles 1 male median lobe, right lateral aspect 2 male median lobe, ventral aspect 3 male right lateral lobe, right lateral aspect 4–6 A. tortus 4 male median lobe, right lateral aspect 5 male median lobe, ventral aspect 6 male right lateral lobe, right lateral aspect.
Sexual dimorphism. Males have the pro-mesotarsomeres I-III distinctly broader than in females with enlarged ventral adhesive setae.
Variation. The few specimens are quite similar to each other in coloration and other features.
This species is known only from southern Suriname (Fig.
The type series was collected from “detrital pools.”
This species is quite unlike other species in the genus. The A. knischi group is characterized by somewhat similarly shaped male median lobes with a fringe of setae along the dorsal margin of each side and many of them have expanded male antennomeres and/or stridulatory devices on the abdomen and metalegs (
This species is named hamatocoles, from Latin hamatus for hooked and coles for penis for the unique shape of the hooked male median lobe in this species (Fig.
Holotype in NZCS, male labeled, “SURINAME: Sipaliwini District 2.005700N, 55.969151W, 337m Sipaliwini Savannah Nature Res. Four Brothers Mts, detrital pools, 31.iii.2017 leg. Short. SR17-0331-01D/ Holotype Agaporomorphus hamatocoles Miller, 2020 [red label with double black line border].” 3 paratypes labeled same as holotype except with “…/Paratype Agaporomorphus hamatocoles Miller, 2020 [blue label with black line border].”
Suriname, Sipaliwini District, Sipaliwini Savannah Nature Reserve, Four Brothers Mountains, 2°00.656'N, 55°59.070'W, 275 m.
This species is in the A. dolichodactylus species group which lacks characteristics of other species groups such as expanded male antennomeres, setae on the dorsal surface of the male median lobe, or stridulatory structures or triangular processes on the abdomen (
Agaporomorphus species, male genitalia. 7–9 A. grandisinuatus 7 male median lobe, right lateral aspect 8 male median lobe, ventral aspect 9 male right lateral lobe, right lateral aspect 10–12 A. mecolobus 10 male median lobe, right lateral aspect 11 male median lobe, ventral aspect 12 male right lateral lobe, right lateral aspect 13–15 A. dolichodactylus 13 male median lobe, right lateral aspect 14 male median lobe, ventral aspect 15 male right lateral lobe, right lateral aspect.
Measurements (N = 3). TL = 3.0–3.2 mm, GW = 1.5 mm, PW = 1.2–1.3 mm, HW = 0.8–0.9 mm, EW = 0.5 mm, FL = 0.7–0.8 mm, FW = 0.2–0.3 mm, TL/GW = 2.0–2.2, HW/EW = 1.6–1.7, FL/FW = 2.9–3.6. Body shape elongate oval, evenly and shallowly curved along lateral margins, curvature continuous between pronotum and elytron.
Coloration . Head, pronotum and elytron orange, similar in coloration throughout dorsal surface. Ventral surface orange, similar in coloration throughout but legs slightly lighter in color.
Sculpture and structure. Head shiny, very finely microreticulate comprised of small isodiametric cells; eyes moderately large (HW/EW = 1.6–1.7). Pronotum shiny, similar microreticulation to head; lateral margin slightly curved, extremely finely beaded, bead absent at anterior angle. Elytron with lateral margin shallowly curved; surface shiny, microreticulation extremely fine, apical half with numerous extremely fine punctures. Prosternum elongate, carinate, prosternal process short, strongly carinate medially. Metaventer and metaventral wings smooth and shiny, with very dense, fine microreticulation. Metacoxa smooth and shiny, similar in microsculpture to metaventer; metacoxal lines distinct, region between metacoxal lines narrow medially; metafemur not unusually broadened (FL/FW = 2.9–3.6).
Male genitalia.
Median lobe complex in shape, asymmetrical; in lateral aspect narrow basally, broadened apically, apically broadly truncate with medial small lobe extending beyond truncation (Fig.
Sexual dimorphism.
Males protarsomeres I–III distinctly broader than in females with four large adhesive setae; females without expansion or adhesive setae. Male mesotarsomeres I–III broader than in females, not as strongly expanded as male protarsomeres I–III, male mesotarsomeres with four large ventral adhesives setae; apex of mesotarsomere V extended into small lobe on posterior margin of apex (Figs
Agaporomorphus species, male mesotarsomere V and mesotarsal claws. 16, 17 A. tortus 16 anterior aspect 17 oblique lateral aspect 18, 19 A. grandisinuatus 18 anterior aspect 19 oblique lateral aspect 20, 21 A. mecolobus 20 anterior aspect 21 oblique lateral aspect 22, 23 A. dolichodactylus 22 anterior aspect 23 oblique lateral aspect.
Variation. There is some minor variation in intensity of coloration of the dorsal surface between specimens but this may be because some specimens are more teneral than others.
This species is only known from southern Suriname (Fig.
The type series was collected from “vegetated pools in savanna.”
This species belongs to the A. dolichodactylus group of Agaporomorphus of
This species is named tortus, Latin for “twisted” for the complex shape of the male median lobe in this species (Figs
Holotype in NZCS, male labeled, “Suriname: Sipaliwini District Sipaliwini Savanna Nature Res. 2°00.656'N, 55°59.070'W, 275 m vegetated pools in savanna 1.iv.2017; leg. A.E.Z. Short SR17-0401-01A/ SEMC1542796 KUNHM-ENT [barcode label]/ Holotype Agaporomorphus tortus Miller, 2020 [red label with double black line border].” 2 paratypes labeled same as holotype except […SEMC1542807…] and […SEMC1516119…] and paratype label, “…Paratype Agaporomorphus tortus Miller, 2020 [blue label with black line border].”
The parsimony analysis resulted in two equally parsimonious trees (L = 17, CI = 82, RI = 92) (Fig.
A. colberti Miller and Wheeler (Fig.
These are the first records of A. colberti from Suriname with previous records from Venezuela (
A. pereirai Guignot (Fig.
Previous records of this species are from Suriname (Cottica River, Moengo, Boven: fig. 25), and Matto-Grosso and Para, Brazil (
Two equally most parsimonious cladograms of Agaporomorphus species derived from parsimony analysis (L = 17, CI = 82, RI = 92): “alt” = alternative equally parsimonious configuration for A. knischi clade. Numbers above hatch marks refer to characters. Numbers below hatch marks refer to character state transformations. Characters mapped using “fast” or “acctran” optimization in WinClada (
Agaporomorphus knischi species group
A. colberti Miller & Wheeler, 2008. Venezuela, Suriname
A. julianeae Hendrich, Apenborn, Burmeister, & Balke, 2013. Peru
A. knischi Zimmermann, 1921. Brazil, Peru, Bolivia
A. sharynae Miller, 2014. Venezuela
A. silvaticus Miller, 2005. Peru
A. tambopatensis Miller, 2005. Peru
Agaporomorphus dolichodactylus species group
A. dolichodactylus Miller, 2001. Brazil, Bolivia
A. grandisinuatus Miller, 2001. Brazil, Peru
A. mecolobus Miller, 2001. Brazil
A. tortus sp. nov. Suriname
Agaporomorphus hamatocoles species group
A. hamatocoles sp. nov. Suriname
Agaporomorphus pereirai species group
A. pereirai Guignot, 1957. Brazil, Suriname
Thanks to Andrew E.Z. Short for collecting and providing access to specimens. Paul Ouboter and Vanessa Kadosoe provided invaluable assistance in Suriname to Short. Portions of this work were funded by NSF grants #DEB–0845984 and #DEB-1353426 (K.B. Miller, PI) and #DEB–0816904 (A.E.Z. Short, PI; K.B. Miller, co–PI). Fieldwork in Suriname was funded by National Geographic Grant #9286-13 (A.E.Z. Short, PI). Finally, thank you to Sandra Brantley for helping with Spanish translation.