Five new species of Trigonopeltastes Burmeister and Schaum from Central America with new country records for other New World Trichiini (Coleoptera, Scarabaeidae, Cetoniinae)

Abstract Five new species of Trigonopeltastes Burmeister and Schaum, 1840 are described: Trigonopeltastes arborfloricola sp. n. from Nicaragua, Trigonopeltastes formidulosus sp. n. from Costa Rica, Trigonopeltastes henryi sp. n. from Costa Rica, Trigonopeltastes mombachoensis sp. n. from Nicaragua, and Trigonopeltastes warneri sp. n. from Belize and Guatemala. An updated key to species of Trigonopeltastes is presented. Trigonopeltastes nigrinus Bates, 1889 and Trigonopeltastes carus Bates, 1889 are placed in synonymy with Trigonopeltastes geometricus Schaum, 1841, syn. n.. The males of Trigonopeltastes aurovelutinus Curoe, 2011 and Trigonopeltastes simplex Bates, 1889 are described for the first time. New country records are given for the following: Giesbertiolus ornatus Howden, 1988: Costa Rica; Paragnorimus sambucus Howden, 1970: Guatemala; Trichiotinus bibens (Fabricius, 1775): Canada; Trigonopeltastes archimedes Schaum, 1841: Guatemala and Costa Rica; Trigonopeltastes frontalis Bates, 1889: Belize, Guatemala, and Honduras; Trigonopeltastes glabellus Howden, 1988: Guatemala; Trigonopeltastes geometricus Schaum, 1841: Honduras; Trigonopeltastes sallaei sallaei Bates, 1889: Guatemala and Honduras; Trigonopeltastes simplex Bates, 1889: Mexico; Trigonopeltastes variabilis Howden, 1968: Honduras.

Etymology. The name arborfloricola approximates "tree flower dweller" in Latin. The name is a noun in apposition. As indicated on the label, the holotype was collected in tree flowers. Distribution (Fig. 56). NICARAGUA -Granada (1): Reserva Privada Silvestre Domitila.
Remarks. This species is fairly distinct in the complete lack of cretaceous markings and orange color pattern on the elytra. Hopefully more specimens will turn up in the lowlands of southern Nicaragua and northern Costa Rica so the intraspecific variation can be described.
Variation. Female allotype (Figs 14-17): length 9.5 mm, width 4.0 mm. The female allotype differs in the following characters. Color: orange band on elytra twice as thick as on holotype (Figs 14-15). Legs dark brown except mesotarsus, metatibia, and metatarsus tan. Pygidium: Surface with smaller basiolateral cretaceous makings. Disc weakly convex with apex not deflexed (Fig. 17). Venter: Sternum without cretaceous markings. Visible abdominal sternite 1 with small, lateral, cretaceous spot; sternites 2-4 without cretaceous markings. Legs: Protibia with 3 distinct teeth, 2 near apex and 1 medial. Paratypes: length 8.5-10.0 mm. Orange band on elytra either thick or thin, regardless of sex; 3 females had greatly expanded, orange coloration covering much of basal half of elytra. Legs vary from dark tan to dark brown with females generally having darker legs than males. Pygidium: Basiolateral cretaceous makings never significantly larger than seen in Figs 12, 17. Venter: Sternum and abdominal sternites of  males with variable-sized cretaceous markings but markings always present in males and never present in females (except on sternite 1).
Etymology. This species is named for the black-and-orange, Halloween-themed coloration of the dorsal surface. The word formidulosus is a Latin for "scary". This name is an adjective in the nominative singular.
Remarks. This species has many similarities to T. geometricus but has consistently different dorsal color pattern and pygidial structure and cretaceous markings. Trigonopeltastes geometricus is variable across it's distribution and within populations but typically has more extensive orange color patterns on the elytra and always has thick lateral bands of cretaceous markings on the pygidium often leaving only a medial strip exposed. Trigonopeltastes formidulosus consistently has a single orange spot on each elytron without further orange lines and markings (except for the occasional thin basal line adjacent to scutellum and three females have more extensive orange coloring covering much of the basal half of the elytra). The apex of the pygidium in males is deflexed at a 90° angle in T. formidulosus while only slightly convex in male T. geometricus. Description of holotype . Male. Length 12.5 mm, width 4.5 mm. Color: head and legs shiny dark metallic green; pronotum and elytra with dull, dark blue, velvety appearance 22); ventral surface shiny black. Head: Surface densely setose medially on disc (except along midline); apex, base, and midline of head moderately to sparsely setose (Fig. 22); clypeus with some longitudinally elongate punctures. Clypeus slightly longer than wide with margins and midline distinctly elevated, apex weakly emarginate. Head without cretaceous markings (Fig. 22). Antenna with 10 antennomeres, club length approximately equal to length of antennomeres 2-7. Maxilla with long, thin brush protruding beyond clypeus in dorsal view. Mentum densely setose, obscuring surface. Pronotum: Surface of disc dull-blue matt with shiny micropunctures (Fig. 22). Marginal bead complete, with complete ring of setose cretaceous markings inside marginal bead. Pronotal disc with more-or-less complete inverted triangle indented into the surface with cretaceous markings, diagonal lines of triangle with row of setae (Fig. 22). Scutellum: Surface setose, without cretaceous markings. Elytra: Surface sparsely setose, matt. Transverse cretaceous band (1 on each side) short, located in basal half of elytral suture (Figs 18-19, near pin but obscured by grease); weak cretaceous markings at apex of suture adjacent to pygidium. Elytral striae well defined with weak indentations between humeral angle and elytral suture. Pygidium: Surface rugose in a circular, fingerprint pattern; densely setose (especially around margins and midline); with thin, inverted U-shaped cretaceous band along basal and lateral surfaces of disc (Fig. 21). Disc strongly, evenly convex. Venter: Sternum with numerous cretaceous markings, densely setose with long setae somewhat obscuring surface. Visible abdominal sternites 2-5 with transverse cretaceous bands thick medially, thin laterally; cretaceous bands setose. Legs: Protibia with 2 teeth near apex (Fig. 19). Mesotibia robust with outer edge bowed outward me-  dially. Tibial spurs acute, unmodified. Tarsi with apicoventral tufts of setae. Holotype with metatarsal legs missing except for 1 metafemur. Parameres: Apically enlarged with a triangular lateral projection (Figs 20,23).
Etymology. This species is named for the late Henry Howden (Ottawa, Canada) as thanks for bringing the female allotype to my attention and to honor his great contributions to our knowledge of Trigonopeltastes and other Trichiini. Temporal data. May (2). Remarks. The male holotype and female allotype are the only known specimens of this new species. The very different dorsal coloration for these specimens is unusual but not unheard of within this genus. I do have some misgivings about placing these two specimens together as one species but decided to take this conservative approach because the structural characters are similar, the size and shape of the two specimens match well, the general cretaceous patterns are congruent, and the two specimens were collected within close proximity to one another. More specimens will need to be examined to understand the color variation within this species and to test my hypothesis that these male and female specimens belong to the same species. . Clypeus slightly wider than long with midline not elevated, apex emarginate. Antenna with 10 antennomeres, club length approximately equal to length of antennomeres 2-7. Maxilla with long, thin brush protruding beyond clypeus in dorsal view. Mentum densely setose, obscuring surface. Pronotum: Surface of disc dull-black with shiny micropunctures (Fig. 31). Marginal bead complete (obscured mediobasally); with complete ring of short, dense, yellow setae and cretaceous markings inside marginal bead. Pronotal disc with more-or-less complete inverted triangle indented into surface; indentation with light cretaceous markings (Fig. 31). Scutellum: Surface with  oblique, cretaceous markings on each side. Elytra: Surface glabrous, matt with shiny micropunctures. Transverse cretaceous band (1 on each side) short, located on basal half of elytral suture. Cretaceous bands (1 on each side) adjacent to lateral edge approximately halfway between base and apex, length approximately ¼ width of elytron (Figs 28-29). Elytral striae 1-4 weakly impressed; remaining striae weakly defined with rows of punctures, not impressed into surface. Orange markings consisting of transverse, basal band; sub-basal, medial spot; 2 longitudinal, medioapical lines . Pygidium: Surface with thick, lateral bands of cretaceous makings that are narrowly joined basally; non-cretaceous area of disc setose, especially near apex (Fig. 32). Disc with distinct, concentric microridges; surface evenly convex with apex slightly deflexed. Venter: Metasternum entirely covered with cretaceous markings and light setae, abdominal sternites with broad, medial cretaceous markings and light setae (Fig. 30). Legs: Protibia with 2 teeth near apex, 1 broad medial tooth (Fig. 29). Mesotibia robust with edges weakly bowed outward medially. Tibial spurs acute, unmodified. Holotype with right mesotarsomeres 2-5 missing. Parameres: Robust with weak lateral notches towards apex (Fig. 33).
Remarks. This species is similar to T. intermedius but can be distinguished by the orange color pattern on the elytra and geographic distribution. Trigonopeltastes intermedius has a solid, transverse, sub-basal, orange line while T. mombachoensis has a sub-basal, orange spot (Figs 28,34). Trigonopeltastes intermedius is known from Mexico and Guatemala and T. mombachoensis is only known from Volcán Mombacho, Nicaragua. Any specimens found of either species in Honduras or northern Nicaragua should be studied to see if these character states are transitional. cept along midline), frons with paired cretaceous markings between eyes (Fig. 41), clypeus with longitudinally elongate punctures. Clypeus about as long as wide with midline distinctly elevated, apex emarginate. Antenna with 10 antennomeres, club length approximately equal to length of antennomeres 2-7. Maxilla with long, thin brush protruding beyond clypeus in dorsal view. Mentum densely setose, setae obscuring surface. Pronotum: Surface of disc dull-black with shiny micropunctures (Fig.  41). Marginal bead complete; with complete ring of setose, cretaceous markings inside marginal bead. Pronotal disc with more-or-less complete inverted triangle indented into the surface with cretaceous markings, cretaceous markings of triangle without setae (Fig. 41). Scutellum: Surface with cretaceous patches in basal corners. Elytra: Surface glabrous, matt. Transverse cretaceous band (1 on each side) short, located on basal half of elytral suture. Cretaceous bands (1 on each side) adjacent to lateral edge approximately halfway between base and apex, length approximately ¼ width of elytron . Elytral striae weakly defined with rows of punctures not indented into surface. Orange markings consisting of transverse, basal band and continuous, oblique T-shaped pattern on each elytron (Figs 38-39). Pygidium: Surface mostly covered with cretaceous makings except for apex and apical 2/3 of medial line (Fig. 40). Disc strongly, evenly convex with apex deflexed, flat. Venter: Sternum mostly covered with cretaceous markings and setae. Visible abdominal sternites 1-5 almost entirely covered with cretaceous markings, with erect setae scattered throughout. Legs: Protibia with 2 teeth near apex (Fig. 39). Mesotibia robust with outer edge weakly bowed outward medially. Tibial spurs acute, unmodified. Holotype with 1 protarsus and 1 metatarsus missing. Parameres: Strongly curved, roughly forming a circle; apex strongly curved inward, enlarged, dorsoventrally flattened (Fig. 42).
Variation. Female allotype (Figs 43-45): length 9.0 mm, width 4.0 mm. The female allotype differs in the following characters: Color: Legs tan to brown. Head: Surface moderately setose with short, obscure setae, head without cretaceous markings (Fig. 45). Pygidium: Surface with thick, inverted U-shaped, cretaceous markings along lateral margins and base; apically and medially without cretaceous markings (Fig. 44). Disc evenly convex without deflexed apex. Venter: Sternum only partially covered with cretaceous markings. Visible abdominal sternites 1-5 without cretaceous markings medially. Legs: Protibia with 3 teeth, 2 near apex and 1 medial. Mesotibia without outer edge bowed outward. Male paratype: length 9.0 mm. Femora and mesotibia orange, protibia half orange and half black longitudinally, metatibia and tarsi black with some dark red. Head: Frons with single cretaceous marking extending between eyes. Clypeus with midline setose, not distinctly elevated. The male paratype is similar in all characters to the holotype.
Etymology. This species is named after Bill Warner (Chandler, Arizona) as thanks for bringing the holotype and allotype specimens to my attention.
Temporal data. May (3). Remark. This species is similar externally to T. sallaei sallaei and T. intermedius but has differences in the elytral coloration pattern. Trigonopeltastes warneri has distinct male parameres (Fig. 42) and is found in the lowlands of eastern Guatemala and Belize, while T. sallaei sallaei and T. intermedius are typically found at mid to high elevation localities.

Notable new distributional records, male descriptions, and new synonymies for New World Trichiini
Giesbertiolus ornatus Howden, 1988 Distribution. This species was previously recorded from Panama (Howden 1988). The specimens detailed below represent a new country record for Costa Rica.

Trichiotinus bibens (Fabricius, 1775)
Distribution. This species was previously recorded from the United States of America (Howden 1968). The specimens detailed below represent a new country record for Canada. These specimens were likely collected about 100-150 years ago. More collecting is needed around London, Ontario and southwestern Ontario to further verify this record. William Saunders (1835-1914) lived in London from 1847-1886 (Bethune 1914) and was a founding member of the Entomological Society of Canada in 1863 and frequent contributor to the early issues of the Canadian Entomologist.

Trigonopeltastes archimedes Schaum, 1841
Distribution. This species was previously recorded from Mexico and El Salvador (Howden 1968). The specimens detailed below represent new country records for Guatemala and Costa Rica.

Trigonopeltastes aurovelutinus Curoe, 2011
Remarks. The male of this species was previously unknown so a description of the key characters is given below. One male specimen was examined labeled "Mex: Guerrero / Acuhuezotla / IX-29-94 Chemsak" (EMEC).
Description of male (Figs 46-51). Color: similar to the description of the females (Curoe 2011;Figs 46-47); dorsal surface black except elytra uniformly orange with darker sutural line. Legs orange with black tarsi and metatibia (Fig. 47). Head: Surface glabrous except for light, setose patches at base of clypeus and apex of frons, without cretaceous markings (Fig. 50). Pronotum: Surface of disc dull-black with shiny micropunctures (Fig. 50); setae scattered across disc but not obscuring surface as seen in the Curoe (2011) figure of the female (perhaps due to abrasion). Marginal bead complete; with thick, scale-like setae. Pronotal disc with more-or-less complete inverted triangle indented into the surface with thick scale-like setae (Fig. 50). Elytra: Orange, glabrous, without cretaceous markings (Figs 46-47). Pygidium: Surface with white, scale-like setae covering disc (but not obscuring surface, perhaps due to abrasion), without cretaceous makings (Fig. 49). Venter: Surface covered with white, scale-like setae. Visible abdominal sternites with erect, white setae scattered medially. Legs: Protibia with 2 teeth near apex (Fig. 47) (female with third medial tooth; Curoe 2011). Metatibia with a distinct, medial protuberance along inner surface (Fig. 46), surface distad to this protuberance smooth with fine striations that may be for stridulation (a key diagnostic character not found in male T. simplex and not apparent in the description and figure of the female T. aurovelutinus). Parameres: Robust with lateral notches towards apex (Figs 48, 51).

Trigonopeltastes frontalis Bates, 1889
Distribution. This species was previously recorded from Mexico (Howden 1968) and El Salvador (Cave 1983). The specimens detailed below represent new country records for Belize, Guatemala, and Honduras.

Remarks. Bates (1889) originally described Trigonopeltastes nigrinus as a variety of
Trigonopeltastes geometricus, and Howden (1968) later gave T. nigrinus full species status. Howden (1968) and Howden and Joly (1998) gave diagnostic characters for each species involving the size, indentation of pronotal triangle, elytral coloration, and pygidial markings but commented that these were based on the examination of "very few specimens" and that further study was needed. In examining longer series of Trigonopeltastes geometricus, I have observed that the diagnostic characters used by Howden (1968) to separate T. geometricus and T. nigrinus break down with a number of individuals exhibiting a blend of the supposed diagnostic characters used. Bates (1889) originally described Trigonopeltastes carus as a distinct species, which Howden (1968) synonymized with T. nigrinus. Specimens that fall under any of the three preceding names all share the diagnostic characters used in the key and have significant lateral portions of the pygidium covered with yellow, cretaceous markings. Since I could find no justification for maintaining T. nigrinus as a separate species, I am here synonymizing this name and T. carus under T. geometricus. Further study involving molecular data is desirable to test the hypothesis that T. geometricus is a single, variable species that is distributed from Mexico to Bolivia.

Trigonopeltastes sallaei sallaei Bates, 1889
Distribution. This species was previously recorded from Mexico, El Salvador, Nicaragua, and Costa Rica (Howden 1968). The specimens detailed below represent new country records for Guatemala and Honduras.

Trigonopeltastes simplex Bates, 1889
Remarks. The male of this species has never been formally described, and so I have included the description of key characters below with sexually dimorphic characters indicated.
Description of male (Figs 52-55). Color: elytra and legs highly variable (see Figs 52-54); dorsal surface black with elytra all black, all dark orange, or with varying degrees of dark orange basally and black apically; each elytron lacking cretaceous markings or with short, transverse, cretaceous band on basal half of elytra suture and/or short cretaceous band adjacent to lateral edge approximately halfway between base and apex. Legs dark orange, black, or a combination of these colors. Head: Surface glabrous, without cretaceous markings. Pronotum: Surface of disc dull-black with shiny micropunctures (surface shiny in females). Marginal bead obsolete laterally, without cretaceous markings inside marginal bead (marginal bead complete and with inside cretaceous markings in females). Pronotal disc with more-or-less complete inverted triangle indented into surface and without cretaceous markings (female has cretaceous markings). Elytra: See above and Figs 52-54 for color variation. Pygidium: Surface without cretaceous makings (present laterally in females), disc with distinct ridges of concentric circles and some setae laterally and apically. Venter: Visible abdominal sternites 1-5 almost covered medially with cretaceous markings (except for small, central triangles); with short, white setae scattered throughout (or long setae just concentrated on sternite 5). Legs: Protibia with 2 teeth near apex (female with third medial tooth). Parameres: Robust with lateral notches towards apex (Fig. 55).
Distribution. This species was previously recorded from Guatemala (Howden 1968). The specimens detailed below represent a new country record for Mexico. The Mexican specimens have some variation in the metathoracic leg characters and setal characters that bear further investigation.
"  Howden, 1968 Distribution. This species was previously recorded from Mexico (Chiapas, San Luis Potosi, and Veracruz), Guatemala, and El Salvador (Howden 1968). The specimens detailed below represent a significant range extension north to the Mexican state of Tamaulipas and a new country record for Honduras.

Updated key to species of Trigonopeltastes
Modified from Howden (1968) and Howden and Joly (1998), and best used with the illustrations in those publications as a reference.
The male of Trigonopeltastes femoratus Howden is unknown and not included in the key.
The females of Trigonopeltastes arborfloricola sp. n., Trigonopeltastes kerleyi Ricchiardi, 2003, andTrigonopeltastes thomasi Howden &Ratcliffe, 1990 are unknown and not included in the key. Pronotum, scutellum, and elytra blue matt (Fig. 19) . Legs bicolored with orange basally and black apically (Fig. 47) Metafemur with short, appressed scales or setae on much of exposed ventral surface (some long setae sometimes at basal posterior edge); pygidium with basal portion of cretaceous markings usually thick and rarely broken medially; elytra with transverse black mark across interval 2 and 3 adjacent to cretaceous marking on midline or intervals 2 and 3 completely orange adjacent to cretaceous marking on midline; elytra along lateral edges with 2 transverse cretaceous lines (basal line sometimes absent especially in individuals with mainly orange coloration on elytra). Mexico to Costa Rica .................Trigonopeltastes sallaei Bates -Metafemur with long, semierect setae (or very elongate flattened scales) on most of exposed ventral surface; pygidium with basal portion of cretaceous markings reduced and sometimes broken medially; elytra with interval 2 orange and interval 3 with black spot or line adjacent to cretaceous marking on midline; elytra along lateral edges with 1 transverse cretaceous line (sometimes absent) (Fig. 28)  Clypeal length approximately equal to width, medial portion of disc with punctures or rugae running longitudinally (e.g., Fig. 16 Howden, 1968-Mexico Trigonopeltastes variabilis Howden, 1968-Mexico, Guatemala, El Salvador, Honduras Trigonopeltastes warneri Smith, 2016-Belize, Guatemala Trigonopeltastes wappesi Howden, 1988 Acknowledgments I thank the late Henry Howden (CMNC) for many interesting discussions on New World Trichiini and for turning over specimens of Trigonopeltastes henryi and T. warneri to me for description. I thank all the curators and collections managers listed in the Materials and Methods section for allowing me to borrow specimens from their collections. Bill Warner (Chandler, Arizona) is acknowledged for allowing me to study the Trigonopeltastes warneri type specimens and Angel Solís (MNCR) is thanked for examining and photographing some of the Trigonopeltastes formidulosus paratypes. Robert Anderson (CMNC) had valuable suggestions that led to the discovery of additional type specimens for two of the new species described. Brett Ratcliffe (UNSM), Enrico Ricchiardi (Torino, Italy), and an anonymous reviewer provided valued reviews of an earlier version of this work.