Theognete chiapaneca Anderson, 2010

Anderson, Robert S., 2010, A taxonomic monograph of the Middle American leaf-litter inhabiting weevil genus Theognete Champion (Coleoptera: Curculionidae; Molytinae; Lymantini) 2458, Zootaxa 2458 (1), pp. 1-127 : 36-41

30. Theognete chiapaneca Anderson , new species ( Figures 98 , 107 , 116 )

Diagnosis. Size. Length, male, 2.75–3.60mm; female, 2.75–4.05mm. Width, male, 1.57–2.25mm; female, 1.46–2.30mm. Pronotum strongly cordate, inflated in males; scales dense, long, erect, recurved, plumose, shaggy, clumped at anterolateral angle and in paramedian tufts; punctures large, dense and deep. Elytra with striae deeply impressed throughout lengths, but not obviously punctate; scales erect, recurved, somewhat concentrated in linear patches along lengths of anterior portions of intervals; posterolateral area with finely punctate crease. Aedeagus as in Fig. 107 .

Geographical distribution. México (Chiapas).

Natural history. Collected from berlese extraction of cloud forest leaf litter at an elevation of 2000m.

Derivation of specific name. Noun in apposition; from the Spanish chiapaneca , meaning a resident of Chiapas.

Material examined. 167♂, 110♀♀ ( BMNH, CMNC, CWOB, ECOS, UNAM, USNM, UVGC) . Holotype ♂ ( UNAM) : MÉXICO: Chiapas. Volcán Tacaná, lower slopes, ca. 4 km. N. Unión Juárez , 2000m, 19.IX.1992, R.S. Anderson, 92-110, cloud forest litter / HOLOTYPE ♂, Theognete chiapaneca sp. nov., R.S. Anderson. Aedeagus extracted. Paratypes: MÉXICO. Chiapas. Volcán Tacaná, lower slopes, ca. 4 km. N. Unión Juárez , 2000m, 19.IX.1992, R.S. Anderson, 92-110, cloud forest litter (18♂, 16♀♀ CMNC, ECOS) . Volcán Tacaná , lower slopes, ca. 4 km. N. Unión Juárez, 2000m, 20.IX.1992, R.S. Anderson, 92-111, cloud forest litter (17♂, 8♀♀ CMNC) . Municipio Angel Albino Corzo, Reserva El Triunfo, Polygono 1, 2050m, 15°39.428'N. 92°48.537'W, 16–21.XI.2001, R.S. Anderson, mixed oak forest, 2001-203 (35♂, 24♀♀ BMNH, CMNC, CWOB, ECOS, USNM, UVGC) ; same data as before except: 2400m, 15°40.16'N 92°48.70'W, cloud forest litter, 2001-204 (47♂, 27♀♀ CMNC) , 2300m, 15°40.16'N 92°48.70'W, 2001-206 (21♂, 13♀♀ CMNC) . Municipio Angel Albino Corzo, 4.3km SE Custepec, Campamento El Quetzal , 18.vii.2007, 15°42ʹN 92°56ʹW, 2160m, M.G.Branstetter #679 (16♂, 15 ♀♀ CMNC) . Municipio Angel Albino Corzo, Reserva Biósfera El Triunfo, Campamento El Quetzal , 17.V.2008, 15.72197° N 92.94163°W, 1850m, R. Anderson, ridge oak forest litter, 2008-010 (1♂, 1♀ CMNC) . Municipio Angel Albino Corzo, 4km SE Custepec, Campamento El Quetzal , 18.V.2008, 15.70318°N 92.93191°W, 2163m, M.G.Branstetter #756 (1♀ CMNC) . Municipio Angel Albino Corzo, 4km SE Custepec, Campamento El Quetzal , 20.V.2008, 15.71119°N 92.93843°W, 2140m, LLAMA Wm-A-03-1-02 (3♂, 1♀ CMNC) . Mpio. Angel Albino Corzo, Reserva Biósfera El Triunfo, Campamento El Quetzal , 1960m ,, 15°.71032 N, 92°.93218 W, 18.V.2008, R.Anderson, mixed, hardwood forest litter 2008-014 (8♂, 4♀♀ CMNC, ECOS) .

Chorological relationships. Sympatric with T. hortenseae , T. confusa , T. galvezi and T. balli .

Theognete tzotzil species group

Recognition. This is the largest species group of Theognete , with 36 species, and consequently the most difficult to characterize. The species are recognized best by the structure of the thoracic sterna. The procoxae tend to be less widely separated than in the similar T. distenta and T. hesmos species groups, and all members have the prosternum in front of the coxae swollen transversely, the middle part of the swelling produced posteriad into a distinct “v”-shape extending between the coxae ( Figs. 8–11 ). This contrasts with T. distenta group members, in which the raised area has a “u”-shaped emargination directed between the coxae ( Figs. 12–15 ), and T. hesmos group members, which lack a raised shelf entirely and at most have an impression or punctures between the coxae ( Figs. 16–18 ). In addition, T. tzotzil group members have the mesosternum flat or sloping slightly anteriad, beset with punctures but no large impressions (28 species) or with some type of deeper pitlike impression (9 species) as in most species of the T. distenta and T. hesmos groups. Lastly, the aedeagus of most T. tzotzil group members is sclerotized across the basal region (25 species).

Diagnosis. Length 2.3–4.8 mm. Cuticle black. Legs moderate in length; femora with small, shallow to moderately deep (4 species) punctures. Pronotal disk weakly (22 species) to strongly (2 species) cordate or subcordate (12 species); medially carinate (1 species) or not; moderately densely, shallowly to moderately deeply punctate, regularly sculptured or with lateral sulcus (9 species), not tuberculate laterodorsally; with vestiture of moderately sparse, fine to coarse, appressed to erect, shallowly to distinctly plumose scales, more or less evenly distributed over disk or concentrated in paramedial and laterodorsal clumps (5 species). Pronotum with flanks flat to subtuberculate anterodorsally. Elytra rounded to elongate-oval, widest in or near middle, smooth (tuberculate in 1 species), striae not evident; with vestiture of fine to coarse, appressed to erect, non-plumose scales scattered evenly over disk, not concentrated in clumps (5 species), to long, distinctly plumose scales; strial punctures not impressed in lateral region of elytra. Posterolateral area of elytra with punctures variously impressed as a series of 3–6 separate to contiguous punctures or with deep pilose crease (3 species). Procoxae narrowly or moderately (7 species) separated. Mesocoxae separated by less than 1.5 x width of coxa. Prosternum swollen transversely in front of coxae, swelling strongly produced posteriad as "V" between coxae, deeply impressed and pit-like posteriorly under swelling, or with swelling not to very slightly produced posteriad between coxae (6 species). Mesosternum flat or slightly sloping anteriad, punctate or impressed to various extent at extreme apex (usually obscured by prosternum), junction with metasternum distinct (26 species) or with median transverse pit-like, usually pilose, impression bounded posteriorly by metasternum (9 species). Metasternum with moderately deep median impression, with distinct, separate small pits or setose punctures laterally behind mesocoxae, which may be subcontiguous with median impression. Abdominal ventrite 5 more or less punctate throughout. Aedeagus short (21 species) to moderate (13 species) in length, apex rounded, not produced to slightly produced (13 species); dorsally not sclerotized (10 species) or sclerotized (24 species) across base.

Comments. This, the largest group of Theognete , encompasses a large degree of subtle interspecific variation, mainly in the pattern and form of the vestiture on the pronotum and elytra. The male genitalia are usually distinctive and unequivocal in distinguishing species. The group is well defined by the distinctive prosternal structure. Thirty of the species in the group occur in Chiapas or countries further south. The group is here divided into five informal groupings based on the form of the dorsal vestiture of the pronotum and elytra. No phylogenetic implications are inferred in these groupings. Group A is characterized by some degree of clumping of the scales on the pronotum and elytra, the scales are usually suberect to erect and dense (often trapping dirt and debris), and the pronotum is generally cordate. Theognete tzotzil ( Fig. 144 ) is typical of this group. Group B has no clumping of scales, the scales are sparser, well-spaced, concentrated in the anterior half of the pronotum and generally linearly arranged on the elytral intervals and suberect or appressed. Most species have a subcordate to weakly cordate pronotum. Theognete campbelli ( Fig. 160 ) is typical. Group C is very similar to B except that the scales of the pronotum and elytra are erect, clavate at the tips, at most only slightly recurved and not obviously plumose. Theognete clavisetosa ( Fig. 136 ) is typical. Group D is very similar to A but with the scales appressed, not clumped on the elytra although condensed into certain areas on the pronotum, the scales are distinctly plumose and the fine hair-like scales of the elytral strial regions are much longer and more robust and very distinct, giving the dorsum of the elytra a shaggy appearance. Theognete huitepec ( Fig.143 ) is typical. Group E is composed of only T. echinata ( Fig. 134 ), which could perhaps also be placed in group C.

Identification of species in this group is best accomplished by geography and then by comparison of sympatric species. The species tend to be highly localized and, aside from the distinctive male genitalia, differ in patterns of pronotal and elytral vestiture, punctation and shape, some of these very subtle. In order to facilitate identifications, three keys for various regional areas are given, one for México (not Chiapas) (6 species), another for México (Chiapas) (21) and another for Guatemala, Honduras and El Salvador (9).

Distribution. Click here for GoogleEarth® maps of T. tzotzil species group distributions.

Key to species of T. tzotzil species group ( México, not Chiapas)

1 Elytra with posterolateral area with single, deep, pilose crease ( Fig. 27 ) .................................................................... 2

1’ Elytra with posterolateral area with series of 3–4 shallow, linearly arranged punctures (as in Fig. 26 ) .................... 3

2 Elytra with basal margin at humeral area with rounded tooth; elytral scales broad, elongate, lanceolate, appressed to suberect, somewhat arched ( Fig. 142 ); aedeagus as in Fig. 177 ............................................................... T. peckorum

2’ Elytra with basal margin at humeral area smooth, lacking tooth; elytral scales fine, elongate, suberect to erect, strongly recurved ( Fig. 164 ); aedeagus as in Fig. 199 .............................................................................. T. ludvigseni

3 Scales of pronotum and elytra very fine, simple, erect and hair-like or bristle-like, not recurved ( Fig. 134 ); aedeagus as in Fig. 170 ............................................................................................................................................... T. echinata

3’ Scales of pronotum and elytra broad, appressed to erect, elongate-oval, acuminate, moderately to strongly recurved, many plumose .............................................................................................................................................................. 4

4 Scales of pronotum and elytral disk evenly distributed, those of elytra linearly arranged along each interval; scales broad, elongate-lanceolate, suberect, not plumose ( Fig. 158 ); aedeagus as in Fig. 193 .................................... T. gaia

4’ Scales of pronotum and elytral disk irregularly distributed, denser in some areas; scales narrow, elongate, suberect to distinctly erect and strongly recurved, many plumose ................................................................................................. 5

5 Scales of elytra suberect, somewhat recurved; scales of elytral strial regions distinct ( Fig. 154 ); aedeagus as in Fig. 189 ............................................................................................................................................................... T. lutulenta

5’ Scales of elytra distinctly erect, strongly recurved; scales of elytral strial regions minute to lacking ( Fig. 147 ); aedeagus as in Fig. 182 ............................................................................................................................... T. howdenorum

Key to species of T. tzotzil species group ( México, Chiapas)

1 Scales of dorsum clumped, forming concentrations (in some specimens dense) variously on the pronotum at anterolateral angles and in paramedian tufts, also in variously placed concentrations on elytra in most specimens ( Figs. 139 , 144, 149, 152 , 162 , 165–166 ) .............................................................................................................................. 2

1’ Scales of dorsum regularly distributed, not clumped, usually sparse, well-spaced, in some specimens concentrated more so in anterior half of pronotum, generally linearly arranged on elytral intervals (as in Figs. 136 , 153 , 161 ) .... 8

2 Elytra elongate, about 1.25x longer than wide ( Figs. 149 , 162 , 166 ) .......................................................................... 3

2’ Elytra rounded, globose, length subequal to width ( Figs. 139 , 144, 152 , 165 ) ........................................................... 5

3 Dorsum with scales simple, erect, not at all arched or recurved at apex, clumped variously in multiple rows over elytral disk ( Fig. 149 ) ....................................................................................................................................... T. montana

3’ Dorsum with scales suberect to erect, plumose, arched or recurved to some extent, not clumped on elytra, although some rows of scales may be denser than others ( Figs. 162–166 ) ............................................................................... 4

4 Aedeagus as in Fig. 197 .................................................................................................................................... T. jonesi

4’ Aedeagus as in Fig. 201 ................................................................................................................................ T. maziaki

5 Elytra with scales clumped in small groups, groups usually distant from each other and wider than one row of scales ( Figs. 152 , 165 ) ............................................................................................................................................................ 6

5’ Elytra with scales clumped in small groups linearly arranged along intervals, usually only as wide as one row of scales (sometimes two) ( Figs. 139 , 144 )....................................................................................................................... 7

6 Elytra with scales of strial regions longer, suberect and arched, giving dorsum a much more shaggy appearance ( Fig. 152 ); aedeagus as in Fig. 187....................................................................................................................... T . scopulus

6’ Elytra with scales of strial regions indistinct ( Fig. 165 ); aedeagus as in Fig. 200 ........................................ T. illuviosa

7 Aedeagus as in Fig. 179 ................................................................................................................................... T. tzotzil

7’ Aedeagus as in Fig. 174 ................................................................................................................................. T. cortesi

8 Scales of elytra erect, clavate at tips, at most only slightly recurved, not obviously plumose ..................................... 9

8’ Scales of elytra appressed to suberect, clavate to acuminate at tips, variously (generally strongly) recurved, distinctly plumose ...................................................................................................................................................................... 11

9 Elytral strial regions with fine, distinct, long, appressed scales; pronotum with scales dense, appressed and distinctly plumose ( Fig. 153 ); aedeagus as in Fig. 188 ............................................................................................ T. backorum

9’ Elytral strial regions lacking scales or with scales very short, very fine and indistinct; pronotum with scales sparse to dense, suberect to erect and not obviously plumose .................................................................................................. 10

10 Pronotum with punctures and vestiture very sparse ( Fig. 136 ); flanks with punctures distinct, subcontiguous, sulci present ( Fig. 136 ); scales of pronotum and elytra straight, recurved only at extreme apex; aedeagus as in Fig. 172 .... .................................................................................................................................................................. T. clavisetosa

10’ Pronotum with punctures and vestiture denser ( Fig. 161 ); flanks with punctures distinct, separate, sulci absent; scales of pronotum and elytra arched; aedeagus as in Fig. 196 .............................................................. T. weadockae

11 Elytra with strial regions with distinct, very long, simple, hair-like scales; pronotum distinctly punctate with scales of disk distinctly plumose ( Fig. 143 ); aedeagus as in Fig. 178.................................................................... T . huitepec

11’ Elytra with strial regions lacking vestiture or, if present, composed of small, short, hair-like scales; pronotum variously punctate to impunctate, with scales of disk simple or plumose ....................................................................... 12

12 Pronotal disk with punctures moderately large to large, dense and deep; basal third similarly punctured to rest of disk; pronotum subcordate ........................................................................................................................................ 13

12’ Pronotal disk with punctures at most small, sparse and shallow; basal third generally impunctate and shining; pronotum subcordate to weakly cordate .............................................................................................................................. 15

13 Elytral intervals with scales short, apically rounded, sparse, not obviously plumose, apices not overlapping ........ 14

13’ Elytral intervals with scales long, apically acuminate, denser, plumose, apices generally overlapping .................. 17

14 Elytra with strial regions with very fine, short, hair-like scales; aedeagus as in Fig. 186 ....................... T. hortenseae

14’ Elytra with strial regions lacking scales; aedeagus as in Fig. 203 .................................................... T. barringerorum

15 Pronotum with punctures moderately large, sparser and shallower; aedeagus as in Fig. 192 ........................... T. balli

15’ Pronotum with punctures large, dense and deep ........................................................................................................ 16

16 Pronotum with impunctate areas in middle and laterally in posterior half; aedeagus as in Fig. 169 ............. T. galvezi

16’ Pronotal disk punctate throughout; aedeagus as in Fig. 190 ......................................................................... T. confusa

17 Pronotal flanks with distinct oblique sulci, no individual punctures evident; aedeagus as in Fig. 181....... T . zapatista

17’ Pronotal flanks with subcontiguous punctures forming faint oblique sulci .............................................................. 18

18 Elytra with strial regions with short, hair-like scales; aedeagus as in Fig. 194 .......................................... T. sylvatica

18’ Elytra with strial regions lacking hair-like scales ..................................................................................................... 19

19 Elytra with scales more robust, apically rounded, well-separated and with apices generally not overlapping; aedeagus as in Fig. 171 ........................................................................................................................................ T. schaubeli

19’ Elytra with scales moderately long, apically tapered or acuminate, more closely spaced and many with apices overlapping ....................................................................................................................................................................... 20

20 Pronotum less cordate ( Fig. 160 ); aedeagus as in Fig. 195......................................................................... T campbelli

20’ Pronotum more cordate ( Fig. 167 ); aedeagus as in Fig. 202 ...................................................................... T. evelynae

Key to species of T. tzotzil species group ( Guatemala, Honduras and El Salvador)

1 Scales of dorsum clumped, variously forming concentrations at anterolateral angles and as paramedian tufts on pronotum, also in variously placed dense concentrations (some more linearly oriented) on elytra ( Figs. 150 , 163 ) ....... 2

1’ Scales of dorsum regularly distributed, not clumped, usually sparse, well-spaced, in some specimens concentrated in anterior half of pronotum, generally variously dense but regularly linearly arranged on elytral intervals .................. 4

2 Elytra with basal margin in humeral region with low, rounded tooth or angulate swelling; elytra elongate, striae with long, fine appressed hairs ( Fig. 163 ); male with middle tibiae distinctly broader along inner margin in middle; aedeagus as in Fig. 198 ..................................................................................................................................... T. coleadesi

2’ Elytra with basal margin in humeral region evenly rounded, lacking swelling; elytra globose, striae with short, fine, appressed hairs or hairs lacking ( Fig. 150 ); male with middle tibiae straight or evenly feebly arcuate along inner margin; aedeagus as in Fig. 185 ................................................................................................................... T. lalannei

3 Elytra with posterolateral region with deep pilose crease (as in Fig. 27 ); pronotum with basal third of disk impunctate, lacking scales and shining; aedeagus as in Fig. 175 ........................................................................... T. trusterae

3’ Elytra with posterolateral region with series of 3–4 separate punctures (as in Fig. 26 ); pronotum variously sculptured but generally basal third with at least some small, fine punctures and/or a few scattered scales ............................... 4

4 Elytral striae with distinct, fine, hair-like scales; pronotum with large, deep punctures .............................. T. lalondei 4’ Elytral striae lacking fine, hair-like scales; pronotum more finely punctate .............................................................. 5

5 Pronotum with punctures very small and fine, widely spaced, indistinct; flanks with punctures distinctly separate; aedeagus as in Fig. 191 ..................................................................................................................... T. lapouimetorum

5’ Pronotum with punctures moderately large and deep, more closely spaced, distinct; flanks with some punctures subcontiguous ................................................................................................................................................................... 6

6 Elytra with scales dense, broadly overlapping along intervals; aedeagus as in Fig. 183 ............................ T. maturini

6’ Elytra with scales less dense, widely separate to very slightly overlapping along intervals ....................................... 7

7 Elytra with scales of disk widely separated along intervals; aedeagus as in Fig. 180 ................................... T. braunae

7’ Elytra with scales of disk very slightly overlapping along intervals ........................................................................... 8

8 Pronotum uniformly distinctly punctate; body length 3.4–4.3mm; aedeagus as in Fig. 173 ...................... T. schusteri

8’ Pronotum with punctures smaller, less distinct, more scattered; body length 3.0– 3.7 mm; aedeagus as in Fig. 175 ..... ...................................................................................................................................................................... T. trusterae

Figures

FIGURES 95–98. Habitus, dorsal view, Theognete bothynometopon species group. 95. T. genieri. 96. T. carinata. 97. T. plumosa. 98. T. chiapaneca.

FIGURES 105–107. Aedeagus, dorsal and lateral view, Theognete bothynometopon species group. 105. T. genieri. 106. T. carinata. 107. T. chiapaneca.

FIGURES 108–116. Abdominal ventrite 5, male, ventral view, Theognete bothynometopon species group. 108. T. jeaneae. 109. T. bothynometopon. 110. T. pragudemi. 111. T. giselae. 112. T. ashei. 113. T. birdi. 114. T. genieri. 115. T. carinata. 116. T. chiapaneca.

FIGURES 8–13. Thoracic sterna, ventral view, Theognete species. 8. T. montana. 9. T. cavei. 10. T. braunae. 11. T. huitepec. 12. T. fossor. 13. T. lalibertei.

FIGURES 14–18. Thoracic sterna, ventral view, Theognete species. 14. T. weiri. 15. T. boaseae. 16. T. hesmos. 17. T. reprela. 18. T. lakarms.

FIGURES 144–154. Habitus, dorsal view, Theognete tzotzil species group. 144. T. tzotzil. 145. T. braunae. 146. T. zapatista. 147. T. howdenorum. 148. T. maturini. 149. T. montana. 150. T. lalannei. 151. T. hortenseae. 152. T. scopulus. 153. T. backorum. 154. T. lutulenta.

FIGURES 155–164. Habitus, dorsal view, Theognete tzotzil species group. 155. T. confusa. 156. T. lapouimetorum. 157. T. balli. 158. T.gaia. 159. T. sylvatica. 160. T. campbelli. 161. T. weadockae. 162. T.jonesi. 163. T. coleadesi. 164. T. ludvigseni.

FIGURES 133–143. Habitus, dorsal view, Theognete tzotzil species group. 133. T. galvezi. 134. T. echinata. 135. T. schaubeli. 136. T. clavisetosa. 137. T. lalondei. 138. T. schusteri. 139. T. cortesi. 140. T. trusterae. 141. T. cavei. 142. T. peckorum. 143. T. huitepec.

FIGURES 25–28. Apex of elytra, lateral view, Theognete species. 25. T. laevis. 26. T. braunae. 27. T. ludvigseni. 28. T. semiopaca.

FIGURES 175–177. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 175. T. trusterae. 176. T. cavei. 177. T. peckorum.

FIGURES 198–199. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 198. T. coleadesi. 199. T. ludvigseni.

FIGURES 169–171. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 169. T. galvezi. 170. T. echinata. 171. T. schaubeli.

FIGURES 193–195. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 193. T. gaia. 194. T. sylvatica. 195. T. campbelli.

FIGURES 187–189. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 187. T. scopulus. 188. T. backorum. 189. T. lutulenta.

FIGURES 181–183. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 181. T. zapatista. 182. T. howdenorum. 183. T. maturini.

FIGURES 165–168. Habitus, dorsal view, Theognete tzotzil species group. 165. T. illuviosa. 166. T. maziaki. 167. T. evelynae. 168. T. barringerorum.

FIGURES 196–197. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 196. T. weadockae. 197. T. jonesi.

FIGURES 200–201. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 200. T. illuviosa. 201. T. maziaki.

FIGURES 178–180. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 178. T. huitepec. 179. T. tzotzil. 180. T. braunae.

FIGURES 172–174. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 172. T. clavisetosa. 173. T. schusteri. 174. T. cortesi.

FIGURES 184–186. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 184. T. montana. 185. T. lalannei. 186. T. hortenseae.

FIGURES 202–203. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 202. T. evelynae. 203. T. barringerorum.

FIGURES 190–192. Aedeagus, dorsal and lateral view, Theognete tzotzil species group. 190. T. confusa. 191. T. lapouimetorum. 192. T. balli.