Enaphalodes Haldeman

Lingafelter, Steven W. & Chemsak, John A., 2002, A New Species Of Enaphalodes Haldeman From Florida (Coleoptera: Cerambycidae) With Review Of Genus, Synonymy, And Key To Species, The Coleopterists Bulletin 56 (4), pp. 569-581 : 570-579

Genus Enaphalodes Haldeman *

Enaphalodes Haldeman 1847: 151 .

Type species: Cerambyx pulverulentus DeGeer 1775 (= Cerambyx atomarius Drury 1773 ). Designated by Linsley 1963:63.

Romaleum White 1855: 309 . Type species: Romaleum operarium White 1855 (= Cerambyx hispicornis Linnaeus 1767 ). By monotypy.

Hypermallus Lacordaire 1869: 302 . Type species: Cerambyx pulverulentus DeGeer 1775 (= Cerambyx atomarius Drury 1773 ). Original designation.

Thersalus Pascoe 1865: 372 . Type species: Phacodes bispinosus Pascoe 1863 (= Cerambyx atomarius Drury 1773 ). By monotypy.

* See Linsley (1963) for additional references.

Diagnosis. Enaphalodes is most closely related to Romulus , Orwellion , Eustromula, Gymnospyra , and Parelaphidion ( Lingafelter 1998) . It differs from Romulus by having more conspicuous pubescence on the elytra and pronotum, moderately spinose elytral apices in most species (truncate to dentiform in Romulus ), more pronounced antennal spination (weak in Romulus ), and smaller size of most specimens. More careful study of Romulus may support its inclusion within Enaphalodes . Enaphalodes , Orwellion , and Eustromula are unique among North American elaphidiines in having the scutellum acutely pointed posteriorly. Enaphalodes differs from Orwellion by lacking differentiated dense, white pubescence posterior to the upper eye lobes (present in Orwellion ). The very short antennomeres of Eustromula (three through five approximately length of pronotum) will differentiate it from Enaphalodes (third through fourth longer than pronotum). Enaphalodes differs from Parelaphidion by its larger size, by having linear (or very weakly, gradually enlarged) metafemora (gradually enlarged in Parelaphidion ), and pronotum inflated at middle and about as wide as base of elytra (not as widely expanded at middle in Parelaphidion ). The length of antennomere three is about two­thirds the length of pronotum in Enaphalodes , which distinguishes it from most Anelaphus and Gymnospyra species (which have antennomere three about half the length of pronotum). Most Anelaphus species have the elytra truncate or dentiform, while Enaphalodes usually have elytral apices moderately to strongly bispinose. Further, Enaphalodes are considerably larger than all Anelaphus .

Key Characters of Enaphalodes . Moderate size, most specimens greater than 20 mm long and 8 mm wide; elytra and pronotum usually covered with dense connected patches of white or yellowish pubescence (uniform in a few species); females with an additional dense coating of short, erect hairs; elytra rarely with sparse pubescence; integument reddish to dark brown; ommatidial facets large; antennomere three of males and most females about two­thirds length of pronotum; mesal antennal spines present; pronotum broadly rounded and inflated, sides at middle projecting to width of base of elytra in males and nearly so in females; prosternal process expanded apically, nearly closing procoxal cavities posteriorly; procoxal cavities closed laterally; pronotum with calli and small impunctate regions; metepisternum very broad anteriorly, strongly narrowing posteriorly; lateral mesosternal projections into mesocoxae absent; femoral apices rounded; metafemoral shape linear or only weakly, gradually enlarged.

Review of Species of Enaphalodes

There are presently nine recognized species of Enaphalodes and all but two have their primary distributions in the United States (Monné 1993; Monné and Giesbert 1993). Enaphalodes atomarius (Drury) ( Fig. 3a ) occurs throughout eastern and central North America into Texas, and south through Mexico and Honduras; E. coronatus (White) ( Fig. 3b ) in Mexico through Costa Rica; E. cortiphagus (Craighead) ( Fig. 3c ) in the northeastern United States west to Arizona; E. hispicornis (Linnaeus) ( Fig. 3d ) throughout the United States, especially the southern half and northern Mexico; E. niveitectus (Schaeffer) ( Fig. 3e ) in the southwestern United States, especially Arizona, and northern Mexico; E. archboldi Lingafelter and Chemsak ( Figs. 1 , 2 , 3f ) in the highlands of south­central Florida, especially the area of the southern Lake Wales Ridge around Archbold Biological Station in Highlands County; E. rufulus (Haldeman) ( Fig. 3g ) in the eastern United States and west to western Texas; E. seminitidus (Horn) ( Fig. 3h ) in the southwestern United States, especially Arizona and California; and E. taeniatus (LeConte) ( Fig. 3i ) in the southwestern United States, especially southern Texas, and northeast Mexico.

Romaleum decipiens Bates 1884 is a New Synonym of Enaphalodes atomarius (Drury 1773) . Examination of the holotype has revealed that there are no characters to distinguish it from the variable E. atomarius (Drury) . Further, Chemsak et al. (1980) demonstrated the distribution of A. atomarius into Zamorano, Honduras, thus enveloping the type locality of Romaleum decipiens Bates ( Paso del Macho , Veracruz, Mexico) well within the revised range of E. atomarius (Drury) . Romaleum cylindricum Knull 1927 is a New Synonym of Enaphalodes cortiphagus ( Craighead 1923) . Examples of both taxa from the type localities has revealed that the only differences are that the Arizona populations have slightly brighter white pubescence and females have a slightly deeper notch (still quite shallow) on the terminal abdominal sternite apex than do the females from Pennsylvania. Because these are both variable characters and no definitive invariable characters have been found to separate the two, we synonymize E. cylindricus Knull.

Key to Species of Enaphalodes

1 Elytra with dense appressed hairs abundant, and conspicuous ( Figs. 3 a­g, i, 5a­c) ------------------------------------------------------------------------------------------------------------------------------------ 2

1' Elytra without appressed hairs and only sparse translucent hairs at apex of elytra and in elytral punctures ( Figs. 3h , 5d ). This species also has weak elytral spines (outer spine often just slightly protuberant); last ventral sternite rounded at apex in females, and with mostly translucent hairs at middle of metasternum and white hairs at sides of metasternum. Length, 24–29 mm- -------------------------------------------------------------------------------------------- E. seminitidus (Horn)

2 Elytra with one to three well­defined areas of dense, appressed pubescence, otherwise ground color not obscured by pubescence ( Fig. 3b, i ) ------------- 3

2' Elytra with pubescence otherwise: either uniform, without patches of dense, appressed pubescence, or with numerous interconnecting, poorly defined appressed patches, giving a mottled appearance ( Fig. 3a , c­g) -- 4

3 Vertex and frons with a very dense, bright white patch of pubescence between the eyes, becoming diffuse and less distinctive toward clypeal margin and on vertex above eyes ( Fig. 3i ); elytra with one transverse patch of white pubescence anterior to middle and a patch on humeri ( Fig. 3i ); apicolateral elytral spine moderate in most specimens; last ventral sternite of female rounded at apex (as in Fig. 4e ). Length, 19–25 mm- ------------------------- -------------------------------------------------------------------------------------------------------------------------- E. taeniatus (LeConte)

3' Vertex and frons with a very dense, bright white patch of pubescence between and above eyes, extending to clypeal margin ( Fig. 3b ); elytra with two well­defined patches of bright white pubescence at apex and around epipleura ( Fig. 3b ); apicolateral elytral spines usually strong; last ventral sternite in females rounded at apex (as in Fig. 4e ). Length, 14–22 mm- ---- ------------------------------------------------------------------------------------------------------------------------------ E. coronatus (White)

4 Elytra not appearing mottled ( Figs. 1–2 , 3d, f ); without patches of appressed pubescence; erect, suberect, and appressed pubescence uniform in density and distribution- --------------------------------------------------------------------------------------------- 5

4' Elytra appearing mottled ( Fig. 3a, c, e, g ); with patches of appressed pubescence; erect, suberect, or appressed pubescence not uniform in density and distribution- ------------------------------------------------------------------------------------------------------------ 6

5 Large species, with pubescence inconspicuous and not heavily obscuring elytral surface ( Fig. 3d ); ventral pubescence mostly translucent with slight golden sheen ( Fig. 5c ); last ventral sternite of females with a deep notch at apex ( Fig. 4d ). Length, 27–40 mm- ----------------- E. hispicornis (Linnaeus)

5' Moderate to large species with conspicuous uniform appressed fulvous pubescence that densely covers elytral surface ( Figs. 1–2 , 3f ); ventral pubescence mostly white and not translucent, distinctly different color from dorsal pubescence ( Fig. 5a ); last ventral sternite of females with a very shallow notch at apex ( Fig. 4b ). Length, 21–30 mm- ------------------------------------------- ------------------------------------------------------------------------------------ E. archboldi Lingafelter and Chemsak

6 Sides of pronotum at middle nearly as wide as base of elytra in males ( Fig. 3a, c, g ); elytra light to dark brown with appressed pubescence extensive but not bright white and not contrasting as intensely with elytral ground color (appearing diffusely mottled) ( Fig. 3a, c, g ); last ventral sternite of female weakly or deeply notched at apex ( Fig. 4a, c, f ) ------------------------------ 7

6' Pronotum with slight widening anterior to middle, but much narrower at widest point than base of elytra ( Fig. 3e ); elytra dark brown with appressed pubescence extensive and conspicuously bright white (distinctly mottled) ( Fig. 3e ); last ventral sternite of female rounded at apex ( Fig. 4e ). Length, 22–27 mm- ---------------------------------------------------------------------- E. niveitectus (Schaeffer)

7 Elytra with white or off­white appressed pubescence and light to dark brown ground color (rarely reddish­brown) ( Fig. 3a, c ); longer erect to suberect setae present (as in Fig. 5a ); last abdominal sternite of females with deep or shallow notch at apex- --------------------------------------------------------------------- 8

7' Elytra with fulvous appressed pubescence and reddish or light brown ground color ( Fig. 3g ); without erect setae (except a few along suture and apical third) ( Fig. 5b ); last abdominal sternite of females with very shallow notch at apex ( Fig. 4f ). Length, 14–26 mm- ----------- E. rufulus (Haldeman)

8 Pubescence of metasternum primarily translucent ( Fig. 5b ); last abdominal sternite of female deeply notched at apex ( Fig. 4a ); elytra with diffuse patches of off­white pubescence ( Fig. 3a ). Length, 19–29 mm- ----------------- ----------------------------------------------------------------------------------------------------------------------------- E. atomarius (Drury)

8' Pubescence of metasternum primarily white and not translucent; last abdominal sternite of female with very shallow notch at apex ( Fig. 4c ); elytra with diffuse patches of white or off­white pubescence ( Fig. 3c ). Length, 16–27 mm- ------------------------------------------------------------------- E. cortiphagus (Craighead)

Figures

Fig. 3. Enaphalodes species, all 1.5 times natural size. a) E. atomarius (Drury) (female); b) E. coronatus (White) (female); c) E. cortiphagus (Craighead) (female); d) E. hispicornis (Linnaeus) (female); e) E. niveitectus (Schaeffer) (female); f) E. archboldi Lingafelter and Chemsak (female); g) E. rufulus (Haldeman) (female); h) E. seminitidus (Horn) (female); i) E. taeniatus (LeConte) (male).

Fig. 1. Enaphalodes archboldi Lingafelter and Chemsak, new species, female dorsal view.

Fig. 2. Enaphalodes archboldi Lingafelter and Chemsak, new species, male dorsal view.

Fig. 5. Lateral view of abdomen and elytron of females. a) Enaphalodes archboldi Lingafelter and Chemsak; b) Enaphalodes rufulus (Haldeman); c) Enaphalodes hispicornis (Linnaeus); d) Enaphalodes seminitidus (Horn).

Fig. 4. Terminal ventral sternite apex of females. a) Enaphalodes atomarius (Drury); b) E. archboldi Lingafelter and Chemsak; c) E. cortiphagus (Craighead); d) E. hispicornis (Linnaeus); e) E. niveitectus (Schaeffer); f) E. rufulus (Haldeman).