Corresponding author: Juan J. Morrone (
Academic editor: M. Alonso-Zarazaga
The phylogenetic relationships of the genera of
Listroderini LeConte, 1876 are one of the largest tribes of
Listroderini were originally assigned to the subfamily
My objective is to analyse the cladistic relationships of the genera of
The studied specimens were provided by the following collections:
American Museum of Natural History, New York, USA.
Amyan MacFadyen, private collection, Coleraine, Northern Ireland.
Alexander Riedel, private collection, Friedberg, Germany.
The Natural History Museum, London, England.
Bernice P. Bishop Museum, Honolulu, USA.
Centro Austral de Investigaciones Científicas, Ushuaia, Argentina.
Carlos Bordón, private collection, Maracay, Venezuela.
Canadian Museum of Nature, Ottawa, Canada.
Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa, Canada.
Charles W. O’Brien private collection, Arizona, USA.
Deutsches Entomologisches Institut, EberswaldeFinow, Germany.
Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil.
Fundación e Instituto Miguel Lillo, San Miguel de Tucumán, Argentina.
Field Museum of Natural History, Illinois, USA.
Guillermo J. Wibmer, private collection, Tallahassee, USA.
Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza, Argentina.
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Santafé de Bogotá, Colombia.
Instituto de la Patagonia, Universidad de Magallanes, Punta Arenas, Chile.
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina.
Museum of Comparative Zoology, Harvard University, Massachusetts, USA.
Museo Nacional de Historia Natural, Santiago, Chile.
Museo de La Plata, La Plata, Argentina.
Museum National d´Histoire Naturelle, Paris, France.
Museo de Zoología “Alfonso L. Herrera”, Facultad de Ciencias, UNAM, Mexico City, Mexico.
New Zealand Arthropod Collection, Auckland, New Zealand.
Staatliches Museum für Tierkunde, Dresden, Germany.
National Museum of Natural History, Washington D.C., USA.
Zoologisk Museum, Copenhagen, Denmark.
Zoologische Museum der Humboldt Universität, Berlin, Germany.
Habitus drawing were made with a camera lucida attached to a stereoscopic microscope. Photographs were taken using a Scanning Electron Microscope at the Facultad de Ciencias, UNAM.
For the present study I examined species of the genera previously recognized for the tribe (
The 58 morphological characters used in the analysis were taken from external structures (53) and male and female genitalia (5). The distribution of character states is shown in the data matrix (
1 Body: length. (0) large to very large (> 15.0 mm); (1) medium-sized (7.1–14.9 mm); (2) small to very small (< 7.0 mm) [additive].
2 Vestiture: scales. (0) present; (1) absent.
3 Vestiture: scale shape. (0) seta-like (
4 Vestiture: setae. (0) present; (1) absent.
5 Rostrum: shape. (0) stout, very short (
6 Rostrum: dorsal carinae. (0) present (
7 Scrobes: shape. (0) long, deep, sharply bordered, reaching eyes; (1) short, ill-defined, broad.
8 Epistome. (0) poorly demarcated; (1) raised.
9 Scrobes: position. (0) dorsolateral to dorsal; (1) lateral.
10 Suprascrobal keels. (0) absent; (1) present.
11 Scrobes: ventral tooth. (0) absent; (1) present (
12 Pterygia. (0) simple, not exposed (
13 Mandibles. (0) with one apical cusp; (1) with two apical cusps.
14 Mandible and pharyngeal processes. (0); short and strong; (1) long and narrow.
15 Mandibles. (0) plurisetose (more than 4 setae); (1) paucisetose (1-4 setae).
16 Maxillary malae: teeth. (0) present; (1) absent.
17 Eyes: shape. (0) subcircular (
18 Eyes: size. (0) large to medium (more than 30 facets); (1) small (10-25 facets); (2) very small (8 or fewer facets) [additive].
19 Eyes: position. (0) lateral (
20 Eyes: convexity. (0) strong; (1) slight; (2) flat [additive].
21 Antennal insertions. (0) distal; (1) at the middle of the rostrum.
22 Scapes: length. (0) long (surpassing posterior margin of eyes when resting in scrobe); (1) medium-sized (reaching eyes when resting in scrobe); (2) short (not reaching anterior margin of eyes when resting in scrobe) [additive].
23 Funicles: segment 1. (0) elongate; (1) globose.
24 Funicles: segments 2. (0) elongate; (1) globose.
25 Funicles: relative lengths of segments 1 and 2. (0) 1 longer than 2 (
26 Funicles: segments 3–6. (0) elongate; (1) globose (
27 Clubs: shape. (0) fusiform; (1) inflated.
28 Pronotum: shape. (0) subcircular; (1) transverse; (2) subtrapezoidal; (3) subquadrate; (4) subclyndrical [non-additive].
29 Pronotum: width. (0) larger than that of elytra; (1) smaller than that of elytra.
30 Pronotum: disc. (0) rugose; (1) smooth, polished.
31 Pronotum: tubercles. (0) absent; (1) present.
32 Postocular lobes. (0) present, well-developed; (1) present, slightly developed; (2) absent [additive].
33 Prosternum. (0) non-excavate; (1) excavate.
34 Metanepisternal sutures. (0) posteriorly fused or obliterated; (1) present, complete.
35 Scutellum. (0) not visible; (1) visible.
36 Elytra: shape. (0) oblong-oval (
37 Elytra. (0) not fused; (1) fused along interelytral suture.
38 Elytral disc. (0) convex; (1) slightly convex; (2) flat [additive].
39 Elytral intervals. (0) convex; (1) flat.
40 Elytral basal margin. (0) not raised; (1) raised, subcarinate.
41 Elytral humeri. (0) rounded; (1) subquadrate.
42 Elytral humeral tubercles. (0) absent; (1) present.
43 Several tubercles on elytral disc. (0) present, small, rounded; (1) absent; (2) present, strong (
44 Series of three tubercles restricted to elytral interval 3. (0) absent; (1) present.
45 Series of declivital tubercles on elytra. (0) absent; (1) present.
46 Carina on elytral apical declivity. (0) absent; (1) present.
47 Anteapical elytral tubercle. (0) absent; (1) present.
48 Elytral apex, female. (0) not produced; (1) produced.
49 Femora: shape. (0) subcylindrical, clavate; (1) dorsoventrally compressed, clavate; (2) subcylindrical, markedly clavate [non-additive].
50 Tibiae: shape. (0) subcylindrical, laterally not expanded; (1) apically expanded.
51 Tibial spurs. (0) present; (1) absent.
52 Tarsomeres 3. (0) bilobed (
53 Ventrites 3 and 4, female. (0) combined shorter than 5; (1) combined longer than 5.
54 Aedeagus, lateral view. (0) robust; (1) slender.
55 Distal gonocoxites. (0) strongly sclerotized; (1) membranous.
56 Styli. (0) well-developed, claw-like; (1) well-developed, finger-like; (2) reduced to a few vibrissae [non-additive].
57 Apodeme of female sternum 8. (0) short (< 3 times longer than plate); (1) long (> 4 times longer than plate).
58 Plate of female sternum 8. (0) developed; (1) reduced.
The cladograms were constructed using software TNT (
Data matrix analysed. Character states of polymorphic taxa are indicated between square brackets.<br/>
|
0000000000000000000000000000000000000000000000000000000000 |
|
1010000010001010100202000100000001000000001000000110000000 |
|
1010000010001010100101000101001011110000001000100010000200 |
|
21?0010010001010010201101100000001000000001000000010001200 |
|
21?0011010011010000101000110000200100000001000010000101210 |
|
2000101010001010100101000101000101100010000110000000001200 |
|
2000101010001010100101000101001101100100000010100000001100 |
|
1000101110101010100100000003000101100010001000000000001[01]00 |
|
2000100011001010110101000104000100100000001000000000011110 |
|
2000100011001010110101000100010101100010001000000000011110 |
|
21?0100011001010020201000114000200100001001000000000001110 |
|
2000200010001010100102000004000101100000001000000000001100 |
|
10[01]0101010001010100101000100000201100000001000100000001[12]00 |
|
2010011010011010100101000101000200100000001010000000001100 |
|
21?0011010011010101201000100000101100000011010002000001100 |
|
20?0011000011010011200010110000200100000001000000000001110 |
|
1000101010001010100101000101000101100000001000100000001100 |
|
2000101010011010100200000004000101100000000000000000001100 |
|
1010200010001010100102001110000101100010001000000000001100 |
|
2011210010001011100112000100000201120010001000000010001201 |
|
1020101010001010100100000101000101100010001000000000011100 |
|
2000200010001010100102000110000101100010001000000000001100 |
|
2000101010001010100100000001001101111000102010100000001100 |
|
21?00110000110100212000101000102001000100010000011000?12?0 |
|
1000001010011010100111000113000201100000001000000010001100 |
|
[12]010101010[01]01010100101000101000101100000001000100000001[01]00 |
|
[12]010[12]00010001010100101001104000101100000001000100000001100 |
|
2010100011001010100101000114000200100000000000000000011110 |
|
2010100010001010100102000104000101110200000000110000001100 |
|
2000100011001010100100000100010201100010001000000000011100 |
|
2010200010001110100102000100000101100010001000000000001100 |
|
2000101000011010010200000114000200000000001000000000001100 |
|
1000210010001010100102001112000101120010001000000000001100 |
|
1000200010001010100102001114000101120010001000000000001100 |
|
0030101010001010110201000100101101011200101010100011001100 |
|
2010100011001010100101000111000101100010000000000000011110 |
|
1030000110001010100201000101001011110000102010100000000000 |
|
2000100010001010100101000101001101110100002001000000001200 |
|
1010200010001010100100000100000101120000001000100000001100 |
|
2030101010001010100100000101001101100100102010100000001100 |
|
2000200010001010100102001114000101120010001000000000001100 |
Some of the characters analysed.
The analysis of the data matrix (
I consider that the results of the analysis with k= 6 are not as extreme as the others and show more clearly the four main clades, which are treated herein as subtribes (
1
2
3
4
Consensus cladograms of the different analyses.
Consensus cladgrma of the cladograms obtained with k=6 with character state changes indicated.
Very small to very large (1.0–22.8 mm); integument reddish brown (black in
Listroderini were formerly considered as related to
Larvae of
1 | Rostrum slender, as long as or longer than pronotum (except shorter than pronotum in some species of |
Palaechthina |
– | Rostrum stout or relatively stout, shorter than pronotum; scrobes usually short, ill-defined, broad; funicular segment 1 longer than 2; associated to terrestrial plants | 2 |
2 | Rostral dorsal carinae usually absent; pterygia auriculate, exposed ( |
Falklandiina |
– | Rostral dorsal carinae present; pterygia simple, not exposed ( |
3 |
3 | Scrobes short, ill-defined, broad, lacking suprascrobal keel; elytra with intervals convex, with anteapical tubercle (except for |
Listroderina |
– | Scrobes long, deep, sharply bordered, reaching eyes, with suprascrobal keel; elytra with intervals usually flat, lacking anteapical tubercle | Macrostyphlina |
Scrobes long, deep, sharply bordered, reaching eyes, with suprascrobal keel; elytra oblong-oval, with intervals usually flat, lacking anteapical tubercle.
This new subtribe, which basically corresponds to the
1 | Postocular lobes present | 2 |
– | Postocular lobes absent | 4 |
2 | Pronotum transverse to strongly transverse | |
– | Pronotum subcircular or subcylindrical | 3 |
3 | Pronotum subcircular with subparallel flanks, disc smooth, polished; metanepisternal sutures present, complete; elytra with intervals flat |
|
– | Pronotum subcylindrical, disc rugose; metanepisternal sutures posteriorly fused or obliterated; elytra with intervals convex | |
4 | Vestiture consisting of subcircular scales and setae; elytra with small, rounded tubercles |
|
– | Vestiture consisting of seta-like scales and setae or only setae; elytra lacking tubercles | 5 |
5 | Vestiture consisting of seta-like scales and setae; eyes large, slightly convex; pronotum disc smooth, polished; basal elytral margin not raised |
|
– | Vestiture consisting of setae only; eyes very small, microphthalmic (8 or fewer facets), flat; pronotum disc rugose; basal elytral margin raised, subcarinate |
|
Small to very small (1.5–4.1 mm); vestiture consisting of seta-like scales and setae; antennal clubs fusiform; pronotum subcylindrical, disc rugose; metanepisternal sutures posteriorly fused or obliterated; elytral intervals convex.
South American Transition Zone (Puna biogeographical province) and Central Chilean and Subantarctic subregions (Andean region), from Peru to Central Chile (
Habitus of representative
Small to very small (3.0–6.6 mm); vestiture consisting of seta-like scales and setae; pronotum subcircular with subparallel flanks, disc smooth, polished; metanepisternal sutures present, complete; elytral intervals flat.
South American Transition Zone (Puna and Coastal Peruvian Desert biogeographical provinces), from Peru to northern Chile (
Very small (1.9–3.3 mm); vestiture consisting of setae only; eyes very small, (8 or fewer facets), flat; postocular lobes absent; basal elytral margin raised, subcarinate.
Species of this genus have been reported as leaf-litter inhabitants (
South American Transition Zone (North Andean Paramo biogeographical province), in Colombia, Ecuador and Peru (
Very small (1.9–3.5 mm); vestiture consisting of subcircular scales and setae; pronotum subclyndrical; metanepisternal sutures posteriorly fused or obliterated; elytra with intervals convex.
South American Transition Zone (North Andean Paramo and Puna biogeographical provinces), from eastern Venezuela to southern Peru (
Small (4.6–6.9 mm); vestiture of seta-like scales and setae; eyes large, slightly convex; pronotum lacking postocular lobes, with disc smooth, polished; elytra with intervals flat.
South American Transition Zone (North Andean Paramo and Puna biogeographical provinces), in Ecuador and Peru (
Small to very small (1.9–6.5 mm); vestiture of subcircular scales and setae; pronotum transverse to strongly transverse; metanepisternal suture present, complete; elytra oblong-oval, with small, rounded tubercles.
Andean region (Subantarctic and Central Chilean subregions) and South American Transition Zone, from southern Argentina, including the Falkland Islands (Islas Malvinas), to Peru (
43;
Rostrum slender, as long as or longer than pronotum (except for some species of
Most of the species of
This subtribe includes the genera
1 | Funicular segment 1 subequal to or shorter than 2 | 2 |
– | Funicular segment 1 longer than 2 | 6 |
2 | Elytra with intervals convex; North and South America | |
– | Elytra with intervals flat; Tristan da Cunha-Gough islands | 3 |
3 | Small to very small (3.7–6.5 mm) |
|
– | Medium-sized to large (7.0–12.0 mm) | 4 |
4 | Vestiture of subcircular scales and setae; pronotum subcircular; elytra oblong-oval |
|
– | Vestiture of seta-like scales and setae; pronotum subtrapezoidal or subclyndrical; elytra elongate-oval | 5 |
5 | Large (11.0–12.0 mm); rostral dorsal carinae absent; pronotum subtrapezoidal |
|
– | Medium-sized (7.0–7.5 mm); rostral dorsal carinae present; pronotum subclyndrical |
|
6 | Scape long (surpassing posterior margin of eye when resting in scrobe); elytra with anteapical tubercle |
|
– | Scape short (not reaching anterior margin of eye when resting in scrobe); elytra lacking anteapical tubercle | 7 |
7 | Vestiture of seta-like scales and setae; Australia and Tristan da Cunha-Gough islands | 8 |
– | Vestiture of subcircular scales ans setae; South America | 9 |
8 | Funicular segments 3-6 elongate; club fusiform; pronotum subclyndrical; elytra with intervals convex; Australia |
|
– | Funicular segments 3-6 globose; club inflated; pronotum subcircular; elytra with intervals flat; Tristan da Cunha-Gough islands |
|
9 | Vestiture of subcircular scales and setae; rostral dorsal carinae present; mandibles long and narrow; antennal insertion distal; postocular lobes slightly developed; elytra oblong-oval; tibiae with spurs | |
– | Vestiture of subcircular scales only; rostral dorsal carinae absent; mandibles robust; antennal insertion at the middle of the rostrum; postocular lobes absent; elytra elongate-oval; tibiae lacking spurs |
Small to very small (2.5–6.0 mm); vestiture of seta-like scales and setae; funicular segments 3-6 elongate; club fusiform; pronotum subclyndrical; elytra with intervals convex.
Australia (
Medium-sized (7.5 mm); vestiture of subcircular scales and setae; pronotum subcircular; elytra oblong-oval.
Tristan da Cunha-Gough islands (
Very small (3.0–3.9 mm); vestiture of subcircular scales only; maxillary mala lacking teeth; antennal insertion at the middle of the rostrum; pronotum subcircular; elytra elongate-oval; tibiae lacking spurs; plate of female sternum 8 reduced.
Bryophytes (
Southern Argentina, including the Falkland Islands (Islas Malvinas), and southern Chile (
Small (4.5 mm); vestiture of seta-like scales and setae; funicular segments 3-6 globose; club inflated; pronotum subcircular; elytra with intervals flat.
Tristan da Cunha-Gough islands (
Very small to medium-sized (1.0–14.0 mm); vestiture of subcircular scales and setae; antennal insertion distal; funicular segment 1 subequal to or shorter than 2; postocular lobes present, well-developed; elytra oblong-oval to elongate-oval, with intervals convex.
Widespread in the Americas, from Canada to Argentina and Chile (
Small (3.8–6.5 mm); vestiture of subcircular scales and setae; mandible and pharyngeal process long and narrow; rostral dorsal carinae present; antennal insertion distal; postocular lobes slightly developed; elytra oblong-oval.
Neotropical region, in Argentina, Bolivia, Chile and Uruguay (
Medium-sized (11.0–12.0 mm); vestiture of seta-like scales and setae; rostral dorsal carinae absent; pronotum subtrapezoidal.
Tristan da Cunha-Gough islands (
Medium-sized (7.0–7.5 mm); vestiture of seta-like scales and setae; rostral dorsal carinae present; pronotum subclyndrical.
Tristan da Cunha-Gough islands (
Small to very small (3.0–6.5 mm); vestiture of subcircular scales and setae; scape long (surpassing posterior margin of eye when resting in scrobe); elytra with anteapical tubercle.
Australia and New Zealand (
Small to very small (3.7–6.5 mm); vestiture of seta-like scales and setae; pronotum subcylindrical.
Tristan da Cunha-Gough islands (
Small to very small (except
This new subtribe, which basically corresponds to the
1 | Scrobes lateral | 2 |
– | Scrobes dorsolateral to dorsal ( |
6 |
2 | Eyes transverse ( |
3 |
– | Eyes subcircular ( |
|
3 | Eyes slightly convex; postocular lobes absent | 4 |
– | Eyes flat; postocular lobes slightly developed | 5 |
4 | Vestiture of setae only; rostrum very short, stout; rostral dorsal carinae absent; scrobes short, ill-defined; eyes dorsal; scape medium-sized (reaching eye when resting in scrobe); funicular segments 3-6 globose; pronotum subcircular; elytra with humeral tubercles; femora subcylindrical, markedly clavate; southern South America | |
– | Vestiture of seta-like scales and setae; rostrum medium-sized, relatively stout; rostral dorsal carinae present; scrobes long, deep, sharply bordered, reaching eyes; eyes lateral; scape long (surpassing posterior margin of eye when resting in scrobe); funicular segments 3-6 elongate; pronotum subcylindrical; elytra lacking humeral tubercles; femora subcylindrical, clavate; New Zealand | |
5 | Very small (2.6–3.5 mm); vestiture of subcircular scales and setae; rostrum lacking dorsal carinae; antennal insertion distal; club fusiform; pronotum transverse; metanepisternal suture posteriorly fused or obliterated; elytra with series of declivital tubercles; tibiae with spurs; southern South America | |
– | Medium-sized (6.0–10.0 mm); vestiture of seta-like scales and setae; rostrum with dorsal carinae; antennal insertion at the middle of the rostrum; club inflated; pronotum subquadrate; metanepisternal suture complete; elytra lacking series of declivital tubercles; tibiae lacking spurs; New Zealand |
|
6 | Rostrum relatively stout, medium-sized, with dorsal carinae; eyes lateral; funicular segment 2 elongate; pronotum subcylindrical; scutellum not visible; New Zealand |
|
– | Rostrum very short, stout, lacking dorsal carinae; eyes dorsal; funicular segment 2 globose; pronotum subcircular; scutellum visible; southern South America | 7 |
7 | Vestiture of seta-like scales and setae; eyes small; club inflated; pronotum with disc rugose; elytra with intervals convex; femora subcylindrical; tibiae subcylindrical | |
– | Vestiture of setae only; eyes very small, microphthalmic; club fusiform; pronotum with disc smooth, polished; elytra with intervals flat; femora dorsoventrally compressed; tibiae apically expanded |
Very small (2.6–3.5 mm); vestiture of subcircular scales and setae; rostrum lacking dorsal carinae; antennal insertion distal; club fusiform; pronotum transverse; metanepisternal suture posteriorly fused or obliterated; elytra with series of declivital tubercles; tibiae with spurs.
Bryophytes (
Andean region (Subantarctic subregion), in southern Argentina, including the Falkland Islands (Islas Malvinas), and southern Chile (
Very small (3.5–4.0 mm); vestiture of setae only; rostrum very short, stout; rostral dorsal carinae absent; scrobes short, ill-defined; eyes dorsal; scape medium-sized (reaching eye when resting in scrobe); funicular segments 3-6 globose; pronotum subcircular; elytra with humeral tubercles; femora subcylindrical, markedly clavate.
Andean region (Subantarctic subregion), in southern Chile (
Small to very small (1.9–6.1 mm); vestiture of seta-like scales and setae; eyes small; club inflated; pronotal disc rugose; elytra with intervals convex.
Andean region (Subantarctic subregion), in southern Argentina, including the Falkland Islands (Islas Malvinas) and southern Chile (
Small to medium-sized (3.5–7.5 mm); vestiture of seta-like scales and setae; rostrum medium-sized, relatively stout; rostral dorsal carinae present; scrobes long, deep, sharply bordered, reaching eyes; eyes lateral; scape long (surpassing posterior margin of eye when resting in scrobe); funicular segments 3-6 elongate; pronotum subcylindrical; elytra lacking humeral tubercles.
New Zealand (
Very small (3.4–3.8 mm); vestiture of setae only; eyes very small, microphthalmic; pronotal disc smooth, polished; femora dorsoventrally compressed; tibiae apically expanded.
The only species of this genus was hypothesized to live in litter or soil (
Andean region (Subantarctic subregion), in the Falkland Islands (Islas Malvinas) (
Habitus of representative
Taxon-area cladogram of the tribe
New Zealand (
Small to very small (2.8–5.0 mm); vestiture of seta-like scales and setae; rostrum relatively stout, medium-sized, with dorsal carinae; eyes lateral; funicular segment 2 elongate; pronotum subcylindrical; scutellum not visible.
New Zealand (
Small (3.9–6.5 mm); vestiture of setae only; eyes subcircular, slightly convex; female elytral apex produced; female ventrites 3 and 4 combined longer than 5.
Andean region (Subantarctic subregion), in southern Chile (
Rostrum relatively stout, medium-sized, shorter than pronotum; scrobes short, ill-defined, broad; funicular segmen 1 longer than 2; elytra usually oblong-oval (subrectangular in
This subtribe, representing the listroderines in the strictest sense, includes the genera
1 | Elytral disc slightly convex to flat | 2 |
– | Elytral disc convex | 6 |
2 | Elytra oblong-oval ( |
3 |
– | Elytra subrectangular ( |
4 |
3 | Vestiture of seta-like scales and setae; scape medium-sized (reaching eye when resting in scrobe) | |
– | Vestiture of scales with finger-like processes and setae; scape long (surpassing posterior margin of eye when resting in scrobe) |
|
4 | Pronotum transverse; elytra with carina on apical declivity, disc slightly convex, lacking anteapical tubercle |
|
– | Pronotum subcircular or subcylindrical; elytra lacking carina on apical declivity, disc flat, with anteapical tubercle |
|
5 | Large (17.5–22.8 mm); mandibles with 3-4 setae; pronotum subcircular, wider than elytra, with tubercles; scutellum not visible; elytra fused along interelytral suture, with series of declivital tubercles; female elytral apex not produced; tibiae lacking spurs; tarsomeres 3 subcylindrical ( |
|
– | Small (4.0–7.0 mm); mandibles with 2 setae; pronotum subcylindrical, narrower than elytra, lacking tubercles; scutellum visible; elytra not fused along interelytral suture, lacking series of declivital tubercles; female elytral apex produced; tibiae with spurs; tarsomeres 3 bilobed ( |
|
6 | Funicular segments 3-6 globose; pronotum lacking tubercles; elytra oblong-oval, not fused along interelytral suture | 7 |
– | Funicular segments 3-6 elongate; pronotum with tubercles; elytra subrectangular, fused along interelytral suture | |
7 | Pronotum transverse or subquadrate; postocular lobes present | 8 |
– | Pronotum subcircular; postocular lobes absent | |
8 | Integument reddish brown; epistome not raised; pronotum transverse | 9 |
– | Integument black; epistome raised; pronotum subquadrate | |
9 | Vestiture of seta-like or lanceolate scales and setae; scrobal ventral tooth absent | 10 |
– | Vestiture of subcircular scales and setae; scrobal ventral tooth usually present | |
10 | Vestiture of seta-like scales and setae | 11 |
– | Vestiture of lanceolate scales and setae |
|
11 | Elytral interval 3 lacking series of three declivital tubercles | |
– | Elytral interval 3 with series of three declivital tubercles |
|
Very small (2.5–3.8 mm); vestiture of seta-like scales and setae; elytra with small, rounded tubercles and series of three tubercles on interval 3.
South American Transition Zone (North Andean Paramo and Puna biogeographical provinces), in Colombia, Ecuador and Peru (
Small (4.0–6.8 mm); vestiture of seta-like scales and setae; scape medium-sized (reaching eye when resting in scrobe); elytra with small, rounded tubercles.
Bolivia, Ecuador and Peru (
Medium-sized (7.3–13.8 mm); integument black; vestiture of seta-like scales and setae; epistome raised; scrobal ventral tooth usually present; pronotum subquadrate.
Andean region (Patagonian subregion), in southern Argentina and southern Chile (
Small to medium-sized (3.7–9.5 mm); vestiture of seta-like or subcircular scales and setae; pronotum subcircular; postocular lobes absent.
Andean region (Subantarctic subregion), in southern Chile and southern
Small to medium-sized (6.0-8.4 mm); vestiture of seta-like scales and setae; pronotum transverse; postocular lobes present.
Andean region (Subantarctic subregion), in southern Chile and southern
Small to medium-sized (5.1–7.5 mm); vestiture of lanceolate scales and setae; postocular lobes present; elytra lacking anteapical tubercle.
Neotropical region and Andean region (Central Chilean subregion), in
Small to medium-sized (5.7–6.8 mm); vestiture of seta-like scales and setae; funicular segments 3-6 elongate; pronotum with tubercles; elytra subrectangular, fused along interelytral suture.
Andean region (Central Chilean subregion) (
Small to medium-sized (3.9–12.5 mm); vestiture of subcircular scales and setae; scrobal ventral tooth usually present.
Andean region (Subantarctic, Central Chilean and Patagonian subregions), South American Transition Zone and Neotropical region, in
Small (4.0–7.0 mm); vestiture of subcircular scales and setae; pronotum subcylindrical, lacking tubercles; scutellum visible; elytra not fused along interelytral suture, lacking series of declivital tubercles; female elytral apex produced; tibiae with spurs.
Australia (
Large to very large (17.5–22.8 mm); vestiture of scales with finger-like processes and setae; mandible with 3-4 setae; pronotum wider than elytra; scutellum not visible; elytra subrectangular, fused along interelytral suture; tibiae lacking spurs; tarsomeres 3 subcylindrical.
Andean region (Subantarctic subregion), in southern Argentina and southern Chile (
Small (5.3–6.6 mm); vestiture of seta-like scales and setae; pronotum transverse; elytra subrectangular, with carina on apical declivity, disc slightly convex, lacking anteapical tubercle.
South American Transition Zone (North Andean Paramo biogeographical province), in Colombia (
Small to very small (2.5–5.3 mm); vestiture of scales with finger-like processes and setae; scape long (surpassing posterior margin of eye when resting in scrobe); pronotum transverse.
Andean region (Central Chilean subregion) (
Ecuador and Peru (
Chile (
The geographical distribution of the genera analysed indicates that
1
2
3
4
By replacing the genera for the areas where they are distributed, a taxon-area cladogram was obtained (
I thank Rolf Oberprieler for stimulating discussions on weevil systematics over the years and for donating specimens of