Phanaeus (Phanaeus) yecoraensis Edmonds, 2004

Phanaeus (Phanaeus) yecoraensis Edmonds View in CoL , new species

Figs. 3–5 View Figs , 8–9 View Figs , 10–14

Type Series. Holotype, male — Mexico: Sonora , Highway 16 , 1.8 road miles [3 km] NW Yécora [28°21'34"N 108°57'06"W], 5,338' [1,640 m] elevation, 6-VIII-2003, R. A. Cunningham and B. D. Streit. GoogleMaps Paratypes (15 males, 14 females)— two males, two females same data as holotype; GoogleMaps one male, four females same data as holotype except 4- VIII-2003; GoogleMaps one male, four females same data as holotype except 5-VIII-2003; three males, three females same data as holotype except 7-VIII-2003; GoogleMaps two males same data as holotype except 18-VII-2003, Barney D. Streit collector; GoogleMaps one female same data as holotype except 5.0 road miles [8 km] NW Yécora [28°20'54"N 108°59'29"W], 5,874' [1,800 m] elevation, 7-VIII-2003; GoogleMaps one male same data as holotype except 1.0 [1.5 km] road mile E Yécora [28°22'20"N 108°54'38"W], 5,147' elevation, 18-VII-2003, Barney D. Streit collector; GoogleMaps one male same data as holotype except 5.0 road miles [8 km] E Yécora [28°23'23"N 108°52'25"W], 5,647' [1,740 m] elevation, 7-VIII-2003; GoogleMaps two males, Sonora, 10 mi [16 km] NW Yécora , 1.3 mi [2 km] S old hwy 16 , Rancho Aguajia , 28-29/ VII 1987, 4,320' [1,330 m], dung trap, S. McCleve; GoogleMaps one male, Sonora, Hwy 16, 20.1 [32 km] mi E Río Yaqui , 3,110 ft [950 m], 26-27/VII 1987, ex dung, S. McCleve; GoogleMaps one male, Sonora, 3.2 mi [5 km] NW Huicoche´ , 5,170 ft [1,590 m], 11–13/VII 1989, ex dung trap, S. McCleve. GoogleMaps (Note: All type specimens bear my determination labels printed in black ink on white paper bordered in red. The holotype and all paratypes collected in 2003 carry secondary labels reading ‘‘Taken in pitfall trap baited with human dung.’’ The female paratype depicted in Fig. 5 View Figs bears an additional label reading ‘‘ Fig. 5 View Figs , Edmonds 2004.’’)

Deposition of Types. Holotype male and three paratypes (1 male; 2 females, including that depicted in Fig. 5 View Figs ) deposited in collection of the Instituto de Ecología (Xalapa, Veracruz — México). Remaining paratypes distributed as follows: National Museum of Natural History, Washington, D.C. (1 male, 1 female); Canadian Museum of Nature, Ottawa (1 male, 1 female); The Natural History Museum, London (1 male, 1 female); private collections of Scott McCleve, Douglas, Arizona (3 males, 1 female); W. D. Edmonds, Marfa, Texas (4 males, 2 females); and Barney D. Streit, Tucson, Arizona, and Richard Cunningham, Chino, California (4 males, 6 females).

Etymology of Species Name. Yécora (place name) þ -ensis (inhabitant of [Latin suffix]). Yécora, Sonora, is the point of reference for the type locality and known distribution of Phanaeus yecoraensis .

Description. Dorsum dark, appearing almost black to unaided eye; magnification (x5) under bright light reveals dark blue, green or bluish-green metallic color on posterior portion of clypeus, paraocular areas, pronotum, elytra and pygidium; pronotum and elytra usually completely colored in male, with black areas of variable size on pronotum and elytra of female; venter dark with green reflections on sterna and legs. Clypeal process quadrate. Protibia tridentate, apical two teeth carinate on outer surface. Posterior pronotal fossae effaced or only weakly indicated. Longer mesotibial spur narrow, not strongly dilated subapically. Elytral interstriae (especially 2–5) flat, broadly shining, finely and evenly punctate (x5; Fig. 9 View Figs ); striae and edges of interstriae shagreened. Male (major): Head bearing large horn, apical one-half abruptly and strongly curved posteroventrally producing hook shape ( Figs. 3–4 View Figs , 11); pronotal disc ( Figs. 3 View Figs , 8 View Figs ) coarsely granulorugose, becoming coarsely punctate posteromedially; sides of pronotum more finely granulorugose than disc, becoming coarsely punctate posterior to lateral fossa; posterior angles of disc strongly rounded and reflexed; disc bordered on each side by narrow sulcus extending from posterior angle to point near anterior margin ( Fig. 8 View Figs , arrow). Male (minor): Pronotum as above except sculpturing finer and punctate areas more extensive, disc lacking lateral sulci; head horn shorter, scarcely curved posteriorly ( Fig. 12 View Figs ) or reduced to small, conical tubercle ( Fig. 13 View Figs ). Female: Head with strongly raised, quadrate process, apex emarginate and dentate laterally ( Figs. 5 View Figs , 14); pronotum evenly granulorugose, becoming punctatorugose to punctate posteriorly; in large individuals, pronotum produced as strong, quadrate lobe, anterior margin carinate; posterior punctation of disc asperate; in smaller individuals, pronotal sculpturing finer, never asperate posteriorly, punctate areas larger and median lobe attenuated, lacking carinate anterior margin. Length [including head]: 15–20 mm; width [at bases of elytra]: 9–12 mm.

Holotype ( Figs. 3–4 View Figs , 8–9 View Figs ):. Large (major) male, length, 19 mm; width, 12 mm. Basal one-half of clypeus metallic green, pronotum greenish-blue, green reflections strongest laterally; elytra dark blue with faint green reflections along anterior and lateral margins; pygidium blue. Posterior pronotal fossae faintly visible. Otherwise as described above.

Specimens examined. 16 males, 14 females.

Discussion. Phanaeus yecoraensis was discovered by Scott McCleve, who very kindly sent me for study four large males collected in dung-baited pitfall traps from three locations west and south of Yécora (see paratype data). At the time (Edmonds 1994) I referred these specimens to Phanaeus flohri Nevinson because of insufficient material, especially females, to make a reliable judgment on their identity. Careful fieldwork by Barney Streit and Richard Cunningham has very recently yielded a series of specimens suitable for careful comparison with P. flohri .

The following modification of the key to the species of the Phanaeus mexicanus group (Edmonds 1994) will distinguish these two species.

1a Elytral interstriae smooth and shiny medially, shagreened laterally, finely punctate (x25); striae dull, shagreened. Sides of pronotum densely punctatorugose around and behind lateral fossae. Pronotal disc of large males flat, posterior angles strongly rounded, entire disc (seen from above) almost heart-shaped. Longer mesotibial spur not strongly dilated subapically. Clypeal process almost quadrate. Dark, appearing almost black to unaided eye (magnification reveals dark blue or green coloration) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2

1b Elytral interstriae evenly shiny, smooth or only very weakly and sparsely punctate (x25). Sides of pronotum granulorugose around and at least some distance behind lateral fossae. Pronotal disc of large male variable but rarely flat. Longer mesotibial spur strongly dilated subapically, especially in female (except scutifer ). Clypeal process evenly rounded. Conspicuously colored, usually very brightly so - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - (couplet 2, Edmonds 1994:61)

2a Cephalic horn of large male robust, distal one-half strongly curved posteroventrally, hook-shaped ( Figs. 3–4 View Figs , 11). Pronotal disc of large male bordered on each side by narrow sulcus ( Fig. 8 View Figs , arrow). Cephalic process of female strongly raised, quadrate, widely bidentate apically ( Figs. 5 View Figs , 14). Elytral interstriae 2–5 broadly shining medially, flat, distinctly punctate (x10; Fig. 9 View Figs ). Northwest Mexico (southeastern Sonora) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Phanaeus (P.) yecoraensis Edmonds , n.sp.

2b Cephalic horn of large male slender, evenly curved posteriorly, never hook-shaped ( Figs. 1–2 View Figs ). Pronotal disc of male lacking lateral sulci ( Fig. 6 View Figs ). Cephalic process of female a low, trituberculate transverse ridge (Fig. 15). Elytral interstriae 2–5 weakly convex, shagreened, only narrowly shiny medially if at all, weakly punctate (x10; Fig. 7 View Figs ). Central Mexico - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Phanaeus (P.) flohri Nevinson

Besides those characters mentioned in the key, P. yecoraensis differs from P. flohri in the following ways: In P. yecoraensis , the posteromedian pronotal fossae are effaced or only weakly indicated, while in P. flohri , they are small but almost always distinct and punctiform; the posterior pronotal angles of large males are relatively more strongly elevated in P. yecoraensis (cf. Figs. 2 View Figs and 4 View Figs ), and the raised posteromedian area of the pronotum of large males is less distinct and more strongly sculptured (the same area appears as a distinctly raised, less punctate area in P. flohri ; cf. Figs. 6 View Figs and 8 View Figs ).

The more strongly elevated posterior pronotal angles of the male produce the false effect of a more flattened prothorax than in P. flohri , which I erroneously observed (Edmonds, 1994). In large females, the median pronotal prominence has a nearly straight, carinate anterior margin and strong lateral angles, while in P. flohri the anterior margin is rounded, not carinate and has weaker lateral angles. Small males and females of P. yecoraensis are best separated from those of P. flohri by the flatter, more conspicuously punctate elytral interstriae (cf. Figs. 7 View Figs and 9 View Figs ) and stronger punctation on posterior portion of pronotum.

The similarities (punctation of the sides of the pronotum; non-dilated mesotibial spurs) I noted in 1994 between P. flohri and the Phanaeus hermes species group extend to P. yecoraensis . Both species share non-dilated mesotibial spurs also with P. scutifer ; and P. yecoraensis and P. scutifer both possess a widely bidentate female cephalic process.

Distribution. The Yécora region lies in the western foothills of the Sierra Madre Occidental near the Sonora — Chihuahua border. Yécora itself (elevation ~ 1,500 m) is located in a small valley between the higher sierras to the east rising to> 2,000 m and supporting pine-oak forests, and to the west rising to ~ 1,700 m and supporting mainly oak-pine-juniper woodlands. Beyond the western ridges, the landscape begins its descent toward the progressively more arid coastal plain of western Mexico. This foothill-coastal plain area supports a rich mixture of scarabaeine faunas produced by dispersal of tropical species from the south and temperate species from the north and east (Mexican Plateau). The known range of Phanaeus yecoraensis lies in this ecogeographic setting. Most collection sites are very near Yécora, but two isolated sites visited by Scott McCleve (pers. comm.) represent the known ecogeographic extremes of this species. The Río Yaqui site (westernmost star symbol in Fig. 10 View Fig ) is about 55 km by road west of Yécora and about 500 m lower in elevation; it supports mesquite scrub woodland with a relictual stand of pines, in stark contrast to the surrounding area. The Huicoché site (~27°19'N 108°37'W; southernmost star symbol in Fig. 10 View Fig ) is about 100 km by air south of Yécora and very near the Chihuahua state boundary. This site also supports pine-oak forest (Scott McCleve, pers. comm); it lies ~ 900 km by air north of the closest known locality for P. flohri (near Lake Chapala, Jalisco).

The main habitat of Phanaeus yecoraensis appears to be oak-pine woodland from about 1,400 m to 1,700 m. Barney Streit and Richard Cunningham (pers. comm.) collected the majority of the type series in human feces-baited pitfall traps just to the west of Yécora, where specimens of P. yecoraensis were captured only during early daylight hours. Their traps at the same location also yielded Deltochilum scabriusculum montanum Howden , Oniticellus rhinocerulus Bates , Dichotomius colonicus (Say) (¼? D. carolinus [L.]), Phanaeus quadridens (Say) , Copris sp. near C. lecontei Matthews , two Onthophagus spp. (one near O. knulli Howden and Cartwright ; one near O. browni Howden and Cartwright ), as well as the geotrupine Geotrupes cavicollis Bates. Also occurring in the Yécora area are the geotrupine Ceratotrupes bolivari Halffter and Martínez and two introduced scarabaeines, Euoniticellus intermedius (Reiche) and Digitonthophagus gazella (F.). The specimen collected by Scott McCleve from the Río Yaqui site west of Yécora is from a lower foothill zone where several other tropical species are collected in Sonora: Phanaeus amithaon Harold , P. furiosus Bates , Coprophanaeus pluto Harold , Dichotomius amplicollis Harold , Canthon indigaceus LeConte , and C. femoralis (Chevrolat) .