C H I A S O G N A T .H I N A E TA X O N O M Y ETHOLOGY OF THE ANDES BIOGEOGRAPHY For the naturalist and the traveller, the American Andes are one of the richest parts of the world f rom a faunistic, geological and botanical point of view – not forgetting the multiplicity of habitats and the history of the peoples who have colonized them. Following in the footsteps of Aimé Bonpland and Alexandre von Humboldt, to mention but them, many naturalists have prospected the Andes and numerous works have been devoted to them, extolling their richness and beauty. In their turn, F. Chalumeau, by profession a writer, and B. Brochier, a pharmacist and biologist, both with a passion for entomology, have devoted some thirteen years to studying a difficult and predominantly Andean subfamily of Coleoptera, consisting of thirty-three representatives in South America, two in Australia, and a fossil one in Europe. The purpose of this book is ambitious. First of all it is to provide a practical tool for the determination of all the surveyed species within an accurate taxonomical f ramework. In order to do this, the authors visited South America many times, with the aim of collecting specimens, and making ecological and faunistic observations. Apart f rom having had the assistance of well-known specialists, they have consulted the collections of the most important national museums (with their precious types) and those of many private individuals. Finally, with the faunistic f ramework created, one of the most important aspects of this work was tackled: the biogeography of the group f rom a worldwide perspective. The reader will no doubt appreciate the complexity of the subject-matter, given the scale of the studied area! Establishing distribution maps may be a relatively easy task, but it is far more difficult to answer questions such as the following: how does one trace the populations across the Andes, and explain the causes of subsequent vicariance in the Chiasognathinae? What conclusions can be drawn? In other words: what are the factors linked to the presence (today or in the past) of the taxa – including the “exotic” species, i.e. those in Australia, and the one found in Europe? Where did all these fauna originate, and how did they survive and multiply? So, apart f rom examining tectonic factors and the geography of the areas covered, and using a floristic approach (and one concerned with other biota), the authors have been obliged to look at the main stages in the history of the earth’s crust, continental drift, and subsequent developments. The result is a work whose language is sometimes complex, but always clear and concise, which is enhanced by plentiful illustrations created by talented professional artists. This book, a classic taxonomic study and an ecological approach to biogeography, is intended for professional and amateur entomologists alike, as well as bio-geographers, but also for all those who are passionate about Nature and wish to discover more about this exceptional region known as the American Andes. They will find here a rich source of information and escapist pleasure.
Summary (Fig. 12 A - B, pages 34 - 35). The Sphaenognathus form a homogenous group within the two subgroups: mandibular subgroup A, with series 4 of the genus; mandibular subgroup B, with the series 1 to 3 of the genus. Nowadays and f rom a morphological perspective, considering the structure of the male mandibles, it would seem that the genus Sphaenognathus represents the most evolved stage of the subfamily. However, species with two mandibular carinae (genera Protognathinus, Chiasognathus, Bomansodus, Chiasognathinus and its subgenera), forming a heterogeneous group, would seem to be less evolved. In the females, the shape of the mandibles is more homogeneous; each of them showing, f rom a dorsal view, the appearance of a half-circle. With the jaws closed, the mouthparts look like an almost circular plateau. One observes a lower carina which is coarsely toothed, an upper carina which is raised and forming the half-circle of the mandible, along with a smaller inner carina and an explanate outer one. As in the males, individual variations are considerable.
Protognathinus
? spielbergi
Australognathus
queenslandicus
Inf. 1A.
grantii beneshi
Chiasognathus
latreillei jousselinii 2 carènes Groupe 1
Sup.
Bomansodus
impubis
peruvianus Chiasognathinus Int. 1B.
gaujoni
Chiasognathinus monguilloni
Sphaenognathinus
? kolbei
mandibularis
Fig. 12A: Mandibular groups and subgroups (1A and 1B).
34
Chiasornithodus
curviceps lindenii metallescens inf. à peine ébauchée 2A.
metallifer série 4 oberon pubescens rotundatus taschenbergi
feisthamelii giganteus 3 carènes Groupe 2
série 3 garleppi higginsi
Sphaenognathus bordoni bellicosus série 2 inf. nette 2B.
armatus nobilis
prionoides spinifer furumii
série 1
alticollis signatus
Fig. 12B: Mandibular groups and subgroups (2A, 2B), and series of Sphaenognathus of the determination keys (1, 2, 3, 4).
35
Map 13: Colombia - climatic areas.
74
Barranquilla 0
200 km
Ca u
ca
Monteria
Cucutá uca
Ara
Magd alena
ue
Puerto Careño
Orén oq
Atrato
Medellín
Bogotá 4
Buenaventura
Tumaco Patia Pasto
are
Guavi
Va upé s Mitú Ya ri
R.
Ne
Caqu
etá
Putumayo
one
Amaz Napo Steppe climate, xerophilous vegetation and rainfalls at midday. Moist climate in the temperate and cold areas. Temperature depends of the altitude (from 18 °C to 20 °C). Moist climate in the cold areas, low páramo (2,500-3,000m) and cloudy forests. High tropical mountain climate. Páramos. Altitude above 3,100 m. Temperature under 10 °C. No vegetation. Very moist climate of equatorial forest, with rainfalls along the year. Moist climate along the year but with short rainfalls times. Pasture climate periodically moist with rainfalls at midday. Desert climate, xerophilous vegetation and rainfalls at midday.
86
gro
Map 14: Colombia - rain flow rates.
74 0
Barranquilla
200 km
a
Monteria
Ca
uc
Cucutá
Medellín
Magd alena
ue
Puerto Careño
Orén oq
Atrato
uca
Ara
Bogotá 4 Buenaventura
are
Guavi
Patia Tumaco
Va upé s
Pasto
Mitú Ya ri
R. N
egr o
Caqu
etá
Putumayo Napo
one
Amaz
less than 500 mm from 500 to 1,000 mm from 1,000 to 2,000 mm from 2,000 to 3,000 mm from 3,000 to 4,000 mm more than 5,000 mm « alto Páramo » area
87
Fig. 40 - 42: Chile, Chiloë Island. Northern valdivian forest (Chilote). Species living there: grantii.
110
Fig. 43 - 46: Chile, Cautín region. Villarica volcano, 1,450 m. Moist valdivian forest. Species living there: grantii.
111
2.3.2.5. The N. dombeyi and N. betuloides evergreen forests. They spread from the Valdivian rain-forest to Tierra del Fuego, occupying the western slopes of the Cordillera and the southern half of Chiloé island. In the northern continental part and in the southern half of Chiloé, the coigüe forest is dominant. These forests reach as far as the 48th parallel, that is to say, far beyond the limits of the Chiasognathus * population. The climate is cool, with an average annual temperature of 8.2 °C and a reduced thermal amplitude. Precipitation decreases from north to south; it ranges from more than 4,000 mm to 1,000-2,000 mm in Tierra del Fuego. First mixed with species from the Valdivian forest which are still present, the coigüe forest quickly becomes predominant, giving way, at around 1,000-1,200 metres altitude, to the lenga forest. Beyond the 48th parallel, the guindo forest (N. betuloides) then appears. 2.3.3. Geonemy of the Chiasognathinae. The data that we have gathered show that the Chiasognathus have only been observed hitherto between 36° south (Longaví) and 45.3° south (Coyhaique), both in Chile and Argentina. With the Andean cordillera separating southern America into two unequal parts, with a distinct geonemy of species, we shall study their distribution in Chile, then in Argentina. A - Chile.
Fig. 52: Chile, Cautín region. Villarica volcano. Moist valdivian forest. Species living there: grantii, impubis. Species likely present: jousselinii, latreillei.
*The extreme distribution limit of grantii is situated near 45.3° (Coyhaique). 116
Chile has been divided into three structural units of southern orientation: coastal range in the west; the Central Valley, and the Andes range, in the east. Natural regions occur in regular succession according to the latitudes and longitudinal zoning. Chiasognathus and Bomansodus are present only in the Valdivian zone. About 1,000 km long (from north to south, between 36° and 48° south), this zone includes the following thirteen administrative regions: Linares, Ñuble, Concepción, Arauco, Bío Bío, Malleco, Cautín, Valdivia, Osorno, Lanquihue, Chiloé, Palena and Aysén. The town of Longaví (36° S., Linares) is probably the northern limit of the range of the Chiasognathinae, with only the jousselinii (topotype of schoenemanni), while Coyhaique (45° S., Aysén) seems to be the southern limit of the expansion of the subfamily, with grantii. Between those two extremes, areas inhabited by chiasognaths are much more numerous between Bío Bío and Llanquihue. Chiasognathus latreillei, jousselinii, grantii, beneshi and Bomansodus impubis are present in the same localities – grantii is the only species on Chiloé. The listed sites are as follows: Linares (Estero de Leiva, Longaví), Ñuble (Chillán, Las Trancas), Concepción (Concepción), Arauco (Contulmo, Lago Lanalhue, Caramavida, Renaico), Bío Bío (Termas Pemehue), Malleco (Angol, Curacautín, Malalcahuello, Lago Jesús María, Las Raíces, Lonquimay, Nahuelbuta, La Fusta, Victoria), Cautín (Lago Caburga, Volcan Villarica, Cautín, Cunco, Cherquenco), Valdivia (Valdivia, Antilhue, Río Bueno, Las Trancas, Río Correntoso), Osorno (Puerto Octay, Puyehue, Antillaca), Llanquihue (Frutillar, Llanquihue, Lago Llanquihue, Lago Chapo, Correntoso, Hornohuinco, Río Blanco), Chiloé (Chiloé island), Palena (Chaitén, Río Michimahuida, Mocopullí, Puerto Cárdenas, Lago Yelcho), Aysén (Aysén, Coyhaique, Puente Zaranda, Río Simpson, Río Cisne Medio, Manihuales).
Fig. 53: Chile, eastern Temuco region, 1,600 m, with view on the Lonquimay volcano. Species likely present: grantii, latreillei, jousselinii, impubis.
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Sphaenognathus oberon Kriesche, 1922,
Sphaenognathus oberon Kriesche, 1922,
C H I A S O G N AT H I N U S G A U J O N I ( O b e r t h ü r, 1 8 8 5 ) (Fig. N° 92 - 98)
Chiasognathus gaujoni Oberthür, 1885. Bull. Soc. Ent. France: 198. Chiasognathinus (Chiasognathinus) (Oberthür) Chalumeau & Brochier, 1995. Derivatio nominis. – As a tribute to the Abbé Gaujon, the first collector of this species. Bibliography: Nonf ried, 1891: 278; Felsche, 1898: 6; Heyne & Taschenberg, 1908: 48, pl. 6, fig. 8; Van Roon, 1910: 4; Kriesche, 1919: 94; Campos, 1921: 50; Didier, 1937: 11; Blackwelder, 1944: 195; Didier & Séguy, 1952: Série A, 28, pl 2, fig 1-8; Didier & Séguy, 1953: Série A, 27: 63 & 71; Benesh, 1960: 150; De Lisle, 1967: 522; Numhauser, 1981: 17; Lacroix, 1984: 7: Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82. Synonymies: [= leoni Lacroix, 1983 (Chalumeau & Brochier, 1995)]. Typical series: Many male specimens. Male lectotype. With the labels: Équateur Loja Abbé Gaujon (black bordered); Chiasognathus Gaujoni R. Oberthür, type Bull. soc. Ent. France 1885. p. CXCVIII (black bordered ); TYPE (red ); MUSÉUM PARIS 200.D.; SYNTYPUS (red ) det. L. Bartolozzi 1988; LECTOTYPE (red ) Chiasognathinus (Oberthür) Dés. F. Chalumeau & B. Brochier ‘92; Sbg CHIASOGNATHINUS Didier & Séguy. – At the MNHN, Paris. Species described from Ecuador. Surveyed localities: Ecuador, region of Loja. Number of specimens examined: 60 M + 3 F. Collections: ANSP (2 M); BMNH (8 M, 1 F); CARN (7 M, 1 F); CAS (3 M); CBOM (6 M, 1 F); CBRO (2 M); FMNH (8 M); IREC (2 M); IRSN (1 M ); MNHUB (6 M); MZ (2 M); SMTD (9 M); UZM (4 M). Diagnosis: Lenght: M: 32-41 mm - F: 28-33 mm. Male: Mandibles black, plunging, imbricate at their apex with, on the upper side, a strong tooth directed towards inside at their base and directed towards the rear end. Under side of the mandible with an acuminate manubrium. Elytra greenish to mauve. Pronotum with violet metallic glints, covered with a yellow ochre pubescence. Legs bicoloure. Female: Same general aspect than the male. Mandibles semi-circular, the upper carina not reaching the base; it finishes at the level of the median tooth of the lower carina. Synonymies • Sphenognathus leoni Lacroix, 1983. Bull. Sci. Nat, 40, p. 7, pl. 2. • Chiasognathinus (Chiasognathinus) leoni (Lacroix) (Chalumeau & Brochier). Derivatio nominis. – As a tribute to Mr Carrère. Bibliography: Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82.
170
Fig. 92 - Chiasognathinus gaujoni Ecuador, Loja
Fig. 93 - Chiasognathinus gaujoni Ecuador, Loja
Fig. 94 - Chiasognathinus gaujoni
Fig. 95 - Chiasognathinus gaujoni
171
Typical series: one specimen. Male holotype. Body length: 30 mm; elytra width: 12 mm; Mandibles length: 5,5 mm – coll. Lacroix. Described from: Perú, Tingo María. X.1969. Allotype not designated. Female unknown. Note: Described by Lacroix from a single specimen, we thought this species was synonym of gaujoni. Indeed, the differences (based on the slender aspect and the coloration of this specimen) are not significant. The examination of a very large series shows, as in all Chiasognathinae, important differences as much in the size as in the colour. In gaujoni, we have observed specimens with elytra either bronze green or reddish brown, and the pronotum with green or violet iridescence. Frequency: Pretty common, but localized.
Fig. 96 - Chiasognathinus gaujoni Ecuador, Loja
172
Fig. 97 - Chiasognathinus gaujoni Ecuador, Loja
Fig. 98 - Chiasognathinus gaujoni Ecuador, Loja
Chiasognathinus gaujoni - x2.8, - x3.1
C H I A S O G N AT H I N U S M O N G U I L L O N I ( L a c r o i x & B o m a n s , 1 9 7 2 ) (Fig. N° 114 - 122)
Sphenognathus monguilloni Lacroix & Bomans, 1972. Bull. Ann. Soc. R. Belg. Ent., 108, page 47, fig. 6. Chiasognathinus (Sphaenognathinus) monguilloni (Lacroix & Bomans) (Chalumeau & Brochier, 1995). Derivatio nominis. – As a tribute to Mr Monguillon. Bibliography: Chalumeau & Brochier, 1995: 20; Krajcik, 2001: 82. Typical series: seven specimens Male holotype. Body length: 30 mm, elytra width: 14,4 mm, mandibles length: 7 mm. With the labels: Sphenognathus monguillonii HOLOTYPE Lacroix J.P. LACROIX det.1971; legue de l ’auteur à MNHN Paris J.P. LACROIX det. 1973 (orange colour bordered ); PERou Mt Carpich 2800 m Tingo Maria Monguillon 1970.; TYPE (red ); Figuré Jla + (stamped signature) – At the MNHN, Paris. With 6 male paratypes: coll. Monguillon, Bomans, Lacroix. Allotype not designated. Species described from Peru, surroundings of Tingo María, Mt Carpish, 2.800 m. Surveyed localities: Peru: regions of Huánuco (Mts Carpish, Tingo María), San Martín (Uchiza Huállagas), Cajamarca (Cajamarca). Number of specimens examined: 93 M + 28 F. Collections: BMNH (1 M, 1 F); CALA (4 M); CARN (4 M); CBOM (4 M, 3 F); CBOR (4 M); CBRO (38 M, 14 F); CM (1 M, 1 F); FMNH (7 M, 2 F); IREC (26 M, 7 F); IRSN (1 M); IZA (3 M); Diagnosis: Lenght: M: 33-58 mm - F: 32-43 mm. Male: Mandibles elongate, slender and plunging; with, at the base of the upper carina, a strong horizontal tooth directed towards inside; on the under side, presence of an acuminate manubrium. Elytra greenish. Lacinia strongly fleshy at its apex. Female: Black. Base of the nasus sinuous. Mandibles broadened and flat on the upper side, with an arc-shaped half-carina more or less raised. Frequency: Common.
184
Fig. 114 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Fig. 115 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Fig. 116 - Chiasognathinus monguilloni
Fig. 117 - Chiasognathinus monguilloni
185
Fig. 118 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Fig. 119 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Fig. 121 - Chiasognathinus monguilloni Peru, Carpish Huánaco
186
Fig. 120 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Fig. 122 - Chiasognathinus monguilloni Peru, Carpish Huánaco
Sphaenognathus monguilloni - x2.25, - x2.9