Zapotecanillus oaxacanus sp. n., by present designation.

The name Zapotecanillus derives from the Zapotecs, the name of the indigenous people living in the territory of Oaxaca during historic times, and the generic name Anillus Jacquelin du Val, the type genus of the subtribe.

The members of this genus are distinguished from the other North and Central American representatives of Anillina by the following combination of characters: frontal area of head flat, without a median tubercle; maxillary palps with palpomere 4 longer than 1/3 of palpomere 3; labium with glossal sclerite with short but distinct paraglossae, and with mentum and submentum fused, without mental-submental suture; pronotum forward of the lateral seta and towards anterior angles with a row of elongate setae; elytra without fixed discal setae and with 8^th and 9^th pores of umbilicate series much closer to each other than the 7^th pore is to the 8^th (i.e. the “geminate” condition). The most distinctive features of the representatives of the new genus, easily distinguishing them from the species of Geocharidius, are the presence of elongate setae at the anterior angles of pronotum forward of the lateral pronotal setae (cf. Fig. 1 versus Fig. 2), a longer maxillary palpomere 4 (cf. Fig. 5 versus Fig. 6), the absence of suture between the mentum and the submentum (cf. Fig. 7 versus Fig. 8), the “geminate” state of the 8^th and 9^th pores of umbilicate series (cf. Fig. 3 versus Fig. 4), the cross-shaped metendoventrite and the truncate intercoxal process between the hind legs (cf. Fig. 13 versus Fig. 14).

Size. SBL range 1.01–1.55 mm.

Habitus. Body form weakly to moderately convex, ovoid or subparallel (Figs 24–31).

Color. Body bicolored (Fig. 24) or monocolorous (Figs 25–31), brunneorufous, rufotestaceous or testaceous, appendages testaceous.

Microsculpture. Dorsal microsculpture of polygonal sculpticells on head, pronotum and elytra. Mesh pattern varies on different body parts. On head, sculpticells transverse, 2-3 times wider than long (Fig. 5). On pronotum, sculpticells longer, 1.5–2 times wider than long. On elytra, sculpticells form mostly isodiametric mesh pattern. Development of microsculpture on pronotum varied among different species.

Luster. Body surface shiny.

Macrosculpture. Body surface sparsely and finely punctate.

Vestiture. Body surface covered with sparse yellow setae of moderate length. Anterior angles of pronotum bear several long setae laterally, which are two times longer than adjacent vestiture (Fig. 1, als).

Fixed setae. Primary head setae include a pair of clypeal (cs), a pair of frontal (fs) and two pairs of supraorbital (ass and pss) setae (Fig. 5). Mentum with three pairs of long primary (medial, paramedial and lateral) setae (Fig. 11, mms, pms, lms). Medial mental setae located on mental tooth, not near its base on mentum (Fig. 7, mms). Submentum with two pairs of long primary setae in two rows (lss1, prss) and 1 additional pair of shorter setae (lss2) located laterally (Fig. 11). Maxilla with long stipetal and palpiferal setae (Fig. 12). Pronotum with two long primary lateral setae (middle, ls, and basal, bs) on each side (Fig. 1). Elytra lack discal setae (Fig. 3), but with scutellar (ed2) and apical (ed8) setae. Last two (8^th and 9^th) pores (eo8 and eo9) of umbilicate series much closer to each other than 7^th (eo7) pore is to 8^th (Fig. 3). Fifth visible sternite of male with two and of female with four setae along the posterior margin.

Head. Anterior margin of clypeus (cl) straight (Fig. 5). Frontal area flat without tubercle (ft) medially near frontoclypeal suture. Fronto-lateral carinae distinct and long.

Eyes. Eyes absent.

Antennae. Submoniliform, 11-segmented, extended to about posterior margin of pronotum. Antennomeres 1 and 2 elongate, of equal length and 1.4–1.5 times longer than antennomere 3, which is only slightly elongate and 1.1–1.2 times longer than antennomere 4. Antennomeres 4 to 10 globose, last antennomere (11) conical and 1.6-1.8 times longer than penultimate antennomere.

Labrum. Labrum (l) transverse with straight, entire anterior margin with six setae apically, increasing in size from the central pair outwards (Fig. 5).

Mandibles. General plan of Bembidion type (Maddison 1993). Right mandible with distinct anterior (art) and posterior retinacular (prt), terebral (tt), premolar (pm) and molar (m) teeth (Fig. 10). Left mandible with distinct terebral (tt), posterior retinacular (prt), premolar (pm) and molar (m) teeth only (Fig. 9).

Maxillae. Maxillary palps (Fig. 12) similar to Bembidion (Maddison 1993) with basal trianguloid cardo, and stipes with dorsal and ventral lobes (dls, vls), dimerous galea (g1, g2), and standard lacinia (lc), with subulate palpomere 4 (mp4). Palpus (Fig. 5) with long 4^th palpomere (mp4), 0.4–0.5 length of palpomere 3 (mp3).

Labium. Labium (Fig. 7) with mental tooth; mentum and submentum fused, without mental-submental suture (ms) and with moderately enlarged lateral mental lobes, which are translucent along the lateral margins (llm). Glossal sclerite (gsc) with short but distinct paraglossae (pg) laterally and with two setae apically. Central area of mental-submental complex with a field of pores and 1-2 pairs of shorter setae additionally (Fig. 11).

Prothorax. Pronotum cordiform, moderately convex, not sinuate (Figs 32–33) or slightly sinuate posteriorly (Figs 34–35). Basal margin of pronotum either straight (Fig. 32), or oblique laterally (Figs 33–34), in one species bisinuate laterally, at posterior angles (Fig. 35). Anterior angles indistinct, broadly rounded. Posterior angles denticulate, without or with 1-2 small denticles in front of the angles. Widths across anterior margin and between posterior angles of approximately equal length (WPa/WPp varies from 0.96 to 1.04 among species).

Scutellum. Externally visible, triangular, with narrowly rounded apex.

Elytra. Elytra of moderate length (LE/SBL from 0.57 to 0.58 among species) without visible interneurs (Fig. 3). Humeri rounded to form right angle with longitudinal axis of body. Basal margination (bm) distinct and long, reaches half the distance between humeral angle and scutellar pore (Fig. 1). Apical half of elytra with shallow but evident subapical sinuation (ss) (Fig. 3).

Hind wings. Absent.

Pterothorax (Fig. 13). Metaventrite (mtv) short, distance between meso- and metacoxae about of the diameter of mesocoxa. Metanepisternum (mte) short, subquadrate, with anterior and outer margins of equal length. Metendoventrite (mes) cross-shaped with long lateral arms.

Legs. Legs of moderate length, not elongate. Prothoracic legs of males variable in structure of tarsomere 1. Typically, 1^st protarsomere markedly dilated apico-laterally with two rows of oval articulo-setae (Stork 1980) on the ventral surface (Figs 20–21). Some species with 1^st tarsomere only slightly dilated and with only one (outer) row of oblong articulo-setae (Fig. 22), and other species with 1^st tarsomere non-dilated and without adhesive vestiture (Fig. 23). Protibiae (Figs 15, 17–19) with antenna cleaner of type B (Hlavac 1971), with both anterior (asr) and posterior (psr) apical setal rows and concave apico-lateral notch (Fig. 18, tbn). Profemora moderately swollen. Mesotibiae with two terminal spurs and tibial brush. Metafemora unmodified, metatibiae with two terminal spurs. Tarsi pentamerous, 5th and 1^st tarsomeres are the longest, 2-4 tarsomeres of equal length on the tarsi of all legs, 1^st tarsomere shorter than combined length of 2–4 tarsomeres (Fig. 15). Tarsal claws simple, untoothed (Fig. 16).

Abdominal ventrites. Five visible abdominal ventrites: 2^nd ventrite longest (Fig. 13), more than 3 times longer than 3^rd or 4^th, 3^rd and 4^th equal in length; the last, 5^th, approximately 1.5 times longer than 4^th. Intercoxal process (ipa) of 2^nd ventrite broad, truncate anteriorly (Fig. 13).

Male genitalia (Figs 36–59). Median lobe of aedeagus anopic, elongate, twisted and slightly arcuate. Internal sac with two groups of copulatory sclerites: dorsal group represented by 2 plates, and ventral group represented by weakly sclerotized fold or folds. Dorsal plate 1 (dp1) in form of an elongate plate, rounded or pointed at basal end, and tapered into a needle-like structure apically. Dorsal plate 2 (dp2) much smaller than plate 1, also needle-attenuated apically, curved and enlarged towards base; coplanarly adjoined to dorsal plate 1 apically in lateral view and divergent from plate 1 basally as a ventrally directed protuberance; can be seen as a separate structure in some species (Figs 54, 57). Ventral sclerites (vsc) of varied shape, dependent on development of sclerotization. Additional spines or scaled membranous fields of internal sac are absent. Parameres typically bisetose, except right paramere of Zapotecanillus pecki 3-setose (Fig. 53). Left paramere large and broad, either evenly tapered to apex (Figs 43, 46) or with short attenuation before setal attachment (Figs 37, 40). Ring sclerite broadly ovate with transverse handle-like extension of varied length and shape (Figs 60–67, hd).

Female genitalia (Figs 68–76). Ovipositor sclerites: Gonocoxite 1 asetose (gc1). Gonocoxite 2 triangular (gc2), 1.6–1.8 times longer than its basal width, slightly to moderately curved, with 2 lateral ensiform (es) and apical nematiform (ns) setae. Laterotergite (lt) with 5–8 setae. Internal genitalia with spermatheca sclerotized, rufous, spherical and ball-shaped in most species, fusiform with a bulb-like enlargement apically in Zapotecanillus kavanaughi (Fig. 75).

The new genus includes eight species: Zapotecanillus oaxacanus sp. n., Zapotecanillus nanus sp. n., Zapotecanillus ixtlanus sp. n., Zapotecanillus iviei sp. n., Zapotecanillus montanus sp. n., Zapotecanillus pecki sp. n., Zapotecanillus kavanaughi sp. n., and Zapotecanillus longinoi sp. n.

The species of this genus are known from three mountain ranges of Mexico: the Sierra Madre de Oaxaca, the Sierra Madre del Sur and the Sierra Madre de Chiapas, within the states of Oaxaca and Chiapas (Fig. 77). This type of distribution best fits Halffter’s (1987) Meso-American Montane Distribution Pattern.

According to the label information, specimens of the new genus were taken from the leaf litter within the 1200–3000m range of altitudes at the Sierra Madre de Oaxaca, and in mesophyll and cloud forests within the 1330–2140m range of altitudes at the Sierra Madre de Chiapas. Collecting dates are May, June, July and August. One specimen from the Sierra Madre de Oaxaca was taken “under bark hardwood”.

The position of Zapotecanillus within the North and Central American Anillina is unclear at present, and awaits molecular data analysis or further discoveries and subsequent morphological analyses of the Middle American anilline taxa. Members of this new genus differ principally from those of the southern stock of Middle American anilline genera (Geocharidius Jeannel, Honduranillus Zaballos, Mexanillus Vigna Taglianti) in having a different arrangement of the last three pores of the umbilicate series and the fused labium, and from geographically proximate Geocharidius and Mexanillus in having distinct paraglossae. They differ from members of the northern stock of North American anilline genera (Anillaspis Casey, Anillinus Casey, Anillodes Jeannel, Micranillodes Jeannel, Serranillus Barr) in lacking fixed discal pores on the elytra.

The key provided below allows distinguishing members of the new genus from those of the other continental North and Middle American anilline genera:

HOLOTYPE, male, in NMNH, point-mounted, labeled: \MEXICO. Oaxaca 18.7mi S Valle Nacional 5200’ 17 Aug.1973\ A.Newton Collector \ Loan from NMNH 2051867\. PARATYPES (8 ex., 4♂2♀ were dissected), labeled same as a holotype, except two specimens, which have an additional label: \Geocharidius n. sp. det. T.L.Erwin 76\ each, where italicized font means handwritten (deposited in CAS, NMNH).

The specific epithet is a Latinized adjective in the masculine form based on Oaxaca, the state of Mexico from which the new species is described.

Mexico, Oaxaca, 18.7mi S from Valle Nacional.

Adults of this new species can be distinguished easily from those of other species of the genus by the following combination of external characters: bicolored and robust appearance, comparatively small head and distinctly transverse pronotum.

Size. Medium-sized for the genus (SBL range 1.30-1.36 mm, mean 1.33±0.049 mm, n=8).

Habitus. Body form (Fig. 24) moderately convex, slightly elongate (WE/SBL 0.41±0.09), head narrow for genus compared to pronotum (WH/WPm 0.69±0.015), pronotum wide compared to elytra (WPm/WE 0.81±0.017).

Color. Body bicolored: head and pronotum brunneorufous, elytra rufotestaceous, appendages testaceous.

Microsculpture. Disc of pronotum with well-developed microsculpture.

Prothorax. Pronotum (Fig. 32) relatively long (LP/LE 0.44±0.006) and markedly transverse (WPm/LP 1.33±0.022), with lateral margins straight and moderately constricted posteriorly (WPm/WPp 1.32±0.034). Basal margin straight. Contour of posterior angles nearly rectangular (100–110°) with 1–2 small denticles in front of the angles.

Elytra (Fig. 3). Convex, not depressed along suture, comparatively wide (WE/LE 0.73±0.013). Margins rounded, slightly divergent in basal half, evenly rounded to apex in apical half, maximal width of elytra at midpoint.

Legs. 1^st male protarsomere only slightly dilated (Fig. 22).

Male genitalia. Median lobe of aedeagus (Fig. 36), with short and transverse apex, broadly rounded at tip. Dorsal plate 1 long, with apical pointed attenuation of moderate length. Dorsal plate 2 joined to plate 1 at its middle ventrally, where it forms a distinct protuberance. Ventral sclerites elongate, with subparallel sides and obliquely stretched from dorsal plates towards ventral margin of median lobe. Right paramere short and moderately wide (Fig. 38). Left paramere with distinct apical constriction (Fig. 37). Ring sclerite with long handle-like extension, widely rounded apically (Fig. 60).

Female genitalia. Gonocoxite 2 rather long, with slightly curved blade and rounded apex (Fig. 68). Laterotergite with 7-8 setae. Spermatheca standard for genus (Fig. 72).

Geographical distribution. This species is known only from the type locality in the Sierra de Juárez Range, a part of the Sierra Madre de Oaxaca, within the high course of the Rio de Valle Nacional (Figs 77 and 94, black quadrangles).

Way of life. Specimens of this species were collected at an altitude of 5200 feet (1600 m).

Relationships. The armature of internal sac of Zapotecanillus oaxacanus males is nearly indistinguishable from that of Zapotecanillus nanus and Zapotecanillus ixtlanus males, described below, clearly suggesting both of them as closest relatives. The former species is sympatric with Zapotecanillus oaxacanus and, based on the same label data, may also be syntopic (i.e., their members may occur together in the same habitat). See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in NMNH, point-mounted, labeled: \ MEX. Oax., 15mi. S. Valle Nacional, 4000’ 21.V.1971 S.Peck Ber.204, leaf litter \ Borrowed Ex. J.M. Campbell \ USNMNH 2051867\. PARATYPES (27 ex., 4♂4♀ were dissected), 4 ex. labeled same as holotype, except one specimen, which has an additional label: \Geocharidius n. sp. det. T.L.Erwin 76\, where italicized font means handwriting.; 12 ex. labeled: MEXICO: Oaxaca, 15mi. S. Valle Nacional, 4000’ 21 May 1971, leaf litter, S. Peck \ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\; 1 ex. labeled: \ MEXICO: Oaxaca 15.1mi S Valle Nacional 4300’ VIII-11-18-1973 \ under bark hardwood A.Newton \ Borrowed ex. MCZ \ Loan from USNMNH 2051867\; 10 ex. labeled: \ MEXICO. Oaxaca 18.7mi S Valle Nacional 5200’ 17 Aug.1973\ A.Newton Collector \ Loan from USNMNH 2051867\ (deposited in CAS, CMNH, NMNH).

The specific epithet is a Latin adjective, nanus, in the masculine form, meaning dwarf, miniature, and refers to the small size of the beetles.

Mexico, Oaxaca, 15mi S from Valle Nacional.

Adults of this new species are distinguished from those of other species of the genus by the combination of small size and brunneorufous color; and males can be further distinguished by the shape of the median lobe (Fig. 39).

Size. Small for genus (SBL range 1.02–1.16 mm, mean 1.10±0.049 mm, n=12).

Habitus. Body form (Fig. 25) moderately convex, slightly elongate (WE/SBL 0.41±0.09), head of normal proportions for genus (WH/WPm 0.75±0.016), pronotum narrow compared to elytra (WPm/WE 0.75±0.028).

Color. Body monocolorous, brunneorufous, appendages testaceous.

Microsculpture. Microlines partially effaced on disc of pronotum.

Prothorax. Pronotum relatively long (LP/LE 0.43±0.020) and moderately transverse (WPm/LP 1.26±0.030), with margins straight and markedly constricted posteriorly (WPm/WPp 1.38±0.044). Basal margin slightly oblique laterally. Posterior angles small, contour of posterior angles obtuse (112–126°) without or with 1 small denticle in front of the angles.

Elytra. Slightly convex, not depressed along suture, comparatively wide (WE/LE 0.72±0.025). Margins rounded, moderately divergent in basal half, evenly rounded to apex in apical third, maximal width of elytra slightly behind the midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally (Fig. 20).

Male genitalia. Median lobe of aedeagus (Fig. 39), with small, slightly elongated apex, angulately rounded at tip. Dorsal plate 1 long, with apical pointed attenuation of moderate length. Dorsal plate 2 joined to plate 1 at its middle ventrally, where it forms a distinct protuberance. Ventral sclerites with sides divergent ventrally, trianguloid in shape. Right paramere short and moderately narrow (Fig. 41). Left paramere with distinct apical constriction (Fig. 40). Ring sclerite with long handle-like extension, pointed apically (Fig. 61).

Female genitalia. Gonocoxite 2 comparatively short, with slightly curved blade and narrowly rounded apex (Fig. 69). Laterotergite with 5-6 setae. Spermatheca standard for genus (Fig. 73).

The species is known from a few localities in the Sierra de Juárez Range, a part of the Sierra Madre de Oaxaca, along the ~5km stretch of the Rio de Valle Nacional (Figs 77 and 94, white triangles).

According to the label data, the elevations of localities range from 4000’ to 5200’ (1200–1600 m).

Aedeagal characters (shape of dorsal plates and left paramere) suggest that Zapotecanillus nanus is most closely related to the sympatric Zapotecanillus oaxacanus. See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in CMNH, point-mounted, labeled: \MEXICO: Oaxaca, 32 miles S Valle Nacional, 7000 ft. 23 May 1971, ex. leaf litter, Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\. PARATYPES (17 ex., 3♂1♀ were dissected), 10 ex. labeled same as a holotype; one male labeled: 8500’, 37mi. S. Valle Nacional, Oax. Mex. V.24.1971 H.Howden \ Borrowed Ex. H.F. Howden \ Loan from USNMNH 2051867\; 3 ex. labeled: \MEXICO: Oaxaca, 37 mi. S Valle Nacional, 8500 ft. 23 May 1971, ex. leaf litter, Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\; 3 ex. labeled: \MEXICO: Oaxaca, 37 mi. S Valle Nacional, oak litter, 25 May 1971, S. Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\ (deposited in CAS, CMNH, NMNH).

The specific epithet is a Latinized adjective in the masculine form based on Ixtlan, the district of the state of Oaxaca, Mexico from which the new species is described.

Mexico, Oaxaca, 32mi. S. Valle Nacional.

Males of this new species can be distinguished from those of other species of the genus by the combination of the large size and shape of the median lobe (Fig. 42).

Size. Large for genus (SBL range 1.32–1.53 mm, mean 1.39±0.069 mm, n=12).

Habitus. Body form (Fig. 26) moderately convex, slightly elongate (WE/SBL 0.41±0.016), head of normal proportions for genus (WH/WPm 0.72±0.013), pronotum narrow compared to elytra (WPm/WE 0.72±0.038).

Color. Body monocolorous, rufobrunneous, appendages testaceous.

Microsculpture. Disc of pronotum with well-developed microsculpture.

Prothorax. Pronotum relatively small (LP/LE 0.39±0.020) and moderately transverse (WPm/LP 1.31±0.039), with margins straight and distinctly constricted posteriorly (WPm/WPp 1.41±0.050). Basal margin oblique laterally. Contour of posterior angles obtuse (111–123°) with a small denticle.

Elytra. Convex, not depressed along suture, of moderate width (WE/LE 0.70±0.023). Margins subparallel at middle, slightly divergent in basal third, evenly rounded to apex in apical third, maximal width of elytra slightly behind midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally.

Male genitalia. Median lobe of aedeagus (Fig. 42) with short semicircular apex. Dorsal plate 1 long, with short apical attenuation. Dorsal plate 2 joined to plate 1 at its middle ventrally, where it forms a distinct protuberance. Ventral sclerites slightly sclerotized, trianguloid in shape. Right paramere short and moderately wide (Fig. 44). Left paramere without apical constriction (Fig. 43). Ring sclerite with short handle-like extension, pointed apically (Fig. 62).

Female genitalia. Spermatheca standard for genus.

The species is known only from the type locality in the Sierra Juárez Range, a part of the Sierra Madre de Oaxaca (Figs 77 and 94, white circle).

Specimens were collected at altitudes 7000-8500’ (2100–2600 m) in oak litter.

Aedeagal characters (shape of the median lobe and dorsal plates) suggest that Zapotecanillus ixtlanus is closely related to Zapotecanillus oaxacanus and Zapotecanillus nanus. See also Fig. 90 for cladistic affinities.

The specific epithet is a Latinized eponym in the genitive case, and is based on the surname of Michael A. Ivie, Associate Professor and Curator of Entomology at the Montana State University, the collector of the type series of the species.

Mexico, Oaxaca, 2 miles S Cerro Pelon.

Adults of this new species can be distinguished from those of other species of the genus by the combination of elongate habitus and comparatively narrow pronotum; and males can be further distinguished by the shape of the copulatory sclerites of the median lobe (Fig. 45).

Size. Large for genus (SBL range 1.34–1.55 mm, mean 1.43 ± 0.066 mm, n=21).

Habitus. Body form (Fig. 27) slightly convex, moderately elongate (WE/SBL 0.39±0.09), head of normal proportions for genus (WH/WPm 0.74±0.013), pronotum narrow compared to elytra (WPm/WE 0.72±0.022).

Color. Body monocolorous, rufotestaceous, appendages testaceous.

Microsculpture. Partially effaced on disc of pronotum.

Prothorax. Pronotum (Fig. 34) relatively short (LP/LE 0.39±0.011) and slightly transverse (WPm/LP 1.22±0.025), with margins slightly sinuate and markedly constricted posteriorly (WPm/WPp 1.42±0.039). Basal margin oblique laterally. Contour of posterior angles obtuse (114–124°) without or with 1 small denticle in front of the angles.

Elytra. Slightly convex, not depressed along suture, rather narrow (WE/LE 0.66±0.019). Margins subparallel at middle, slightly divergent in basal forth, evenly rounded to apex in apical forth, maximal width of elytra at midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally (Fig. 21).

Male genitalia. Median lobe of aedeagus (Fig. 45), with small, slightly elongated apex, angulately rounded at tip. Dorsal plate 1 long, with long apical attenuation. Dorsal plate 2 joined to plate 1 at its middle ventrally, where it forms a distinct protuberance. Ventral sclerites weakly sclerotized. Right paramere rather long and narrow (Fig. 47). Left paramere without apical constriction (Fig. 46). Ring sclerite with short handle, which is widely rounded apically (Fig. 63).

Female genitalia. Spermatheca (Fig. 74) standard for genus.

The species is known only from the type locality in the Sierra Juárez Range, a part of the Sierra Madre de Oaxaca (Figs 77 and 94 black star).

According to the label data (elevation ranges 2600–2700m), these beetles inhabit the pine-oak forest zone of the Sierra Madre de Oaxaca.

Externally, adults of Zapotecanillus iviei are similar to those of Zapotecanillus kavanaughi, Zapotecanillus pecki and Zapotecanillus montanus, described below, but the armature of the internal sac of the median lobe suggests closer relatedness to Zapotecanillus oaxacanus, Zapotecanillus nanus and Zapotecanillus ixtlanus. See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in CMNC, point-mounted, labeled: \MEX. Oax. 14km N SanJuan del Estado 2600m. 4-VIII.1986 H. & A. Howden\ berlese\ CMNC\. PARATYPES (8 ex., 2♂2♀ were dissected), labeled same as a holotype (deposited in CAS, CMNC).

The specific epithet is a Latinized eponym in the genitive case, and is based on the surname of David H. Kavanaugh, Senior Curator of the Entomology Department of the California Academy of Sciences, whose enthusiastic efforts in locating and borrowing the material for the current investigation were so magnanimous and productive.

Mexico, Oaxaca, 14 km N San Juan del Estado.

Adults of this new species are distinguished from those of other species of the genus by the combination of elongate habitus and comparatively narrow pronotum; and males can be further distinguished by the shape of median lobe (Fig. 54).

Size. Medium-sized for genus (SBL range 1.25–1.42 mm, mean 1.34±0.055 mm, n=9).

Habitus. Body form (Fig. 28) slightly convex, moderately elongate (WE/SBL 0.38±0.13), head of normal proportions for genus (WH/WPm 0.77±0.014), pronotum narrow compared to elytra (WPm/WE 0.74±0.020).

Color. Body monocolorous, rufotestaceous, appendages testaceous.

Microsculpture. Partially effaced on disc of pronotum.

Prothorax. Pronotum (Fig. 33) relatively short (LP/LE 0.40±0.009) and slightly transverse (WPm/LP 1.23±0.031), with margins rectilinear and distinctly constricted posteriorly (WPm/WPp 1.36±0.039). Basal margin oblique laterally. Contour of posterior angles obtuse (116–122°) with 1–2 small denticles in front of the angles.

Elytra. Slightly convex, not depressed along suture, rather narrow (WE/LE 0.66±0.023). Margins almost subparallel, slightly divergent in basal half, evenly rounded to apex in apical third, maximal width of elytra posterior to midpoint.

Legs. 1^st male protarsomere not dilated, without adhesive vestiture (Fig. 23).

Male genitalia. Median lobe of aedeagus (Fig. 54), with very narrow, elongate apex. Dorsal plate 1 short, pointed basally, with apical attenuation of moderate length. Dorsal sclerite 2 in a form of a separate structure, crosses plate 1 at apical third. Ventral sclerites slightly sclerotized. Right paramere rather long and moderately wide (Fig. 56). Left paramere without apical constriction (Fig. 55). Ring sclerite with long handle-like extension, widely rounded apically (Fig. 66).

Female genitalia. Gonocoxite 2 comparatively short, with moderately curved blade and rounded apex (Fig. 70). Laterotergite with 5-6 setae. Spermatheca atypical for genus (Fig. 75).

The species is known only from the type locality in the Sierra Aloapaneca Range, a part of the Sierra Madre de Oaxaca (Figs 77 and 94, black circle).

All beetles were collected at an elevation of 2600 m.

Externally, adults of Zapotecanillus kavanaughi are similar to those of Zapotecanillus iviei, Zapotecanillus pecki and Zapotecanillus montanus, described below, but males and females differ from those of these species in features of the median lobe and shape of the spermatheca, respectively. See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in CMNH, point-mounted, labeled: \ MEXICO, Oaxaca, 52miles N of Oaxaca, Ber.202, sink litter, 17 May 1971 S.B.Peck collector\ CMNH \. PARATYPES (14 ex., 3♂4♀ were dissected), 6 ex. labeled same as a holotype; one female labeled: \ MEX. Oax., 52mi. N Oaxaca, 9500’ 17.V.71 S.Peck Ber.202, leaf lit.\ Anillinus (handwriting)\; 7 ex. labeled: \MEXICO: Oaxaca, 52miles N Oaxaca, 9500 ft., 25 May 1971, ex. litter in sinkhole, S.Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\ (deposited in CAS, CMNH).

The specific epithet is a Latin adjective from mons (= mountain), in the masculine form, meaning mountain-dwelling, and refers to the altitudinal data of the species locality.

Mexico, Oaxaca, 52 miles N of Oaxaca.

Recognition. Males of this new species are distinguished from those of other species of the genus by the combination of elongate habitus and shape of the median lobe (Fig. 48).

Size. Medium-sized for genus (SBL range 1.29–1.40 mm, mean 1.35±0.037 mm, n=7).

Habitus. Body form (Fig. 29) slightly convex, moderately elongate (WE/SBL 0.40±0.10), head of normal proportions for genus (WH/WPm 0.75±0.041), pronotum narrow compared to elytra (WPm/WE 0.70±0.012).

Color. Body monocolorous, rufotestaceous, appendages testaceous.

Microsculpture. Partially effaced on disc of pronotum.

Prothorax. Pronotum relatively short (LP/LE 0.38±0.009) and moderately transverse (WPm/LP 1.26±0.028), with margins rectilinear and markedly constricted posteriorly (WPm/WPp 1.40±0.028). Basal margin oblique laterally. Contour of posterior angles obtuse (115–125°) with 1–2 small denticles in front of the angles.

Elytra. Convex, not depressed along suture, of moderate width (WE/LE 0.69±0.016). Margins subparallel at middle, slightly divergent in basal third, evenly rounded to apex in apical third, maximal width of elytra at midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally.

Male genitalia. Median lobe of aedeagus (Fig. 48) with slightly elongate apex, angulately rounded at tip. Dorsal plate 1 short, pointed apically and basally. Dorsal plate 2 joined to plate 1 at its middle ventrally, where it forms a distinct protuberance. Ventral sclerites with pronounced sclerotization ventrally. Right paramere short and moderately wide (Fig. 50). Left paramere without apical constriction (Fig. 49). Ring sclerite with handle conically rounded apically (Fig. 65).

Female genitalia. Spermatheca standard for genus.

The species is known only from the type locality in the Sierra Juárez Range, a part of the Sierra Madre de Oaxaca (Figs 77 and 94, white quadrangle).

Specimens of this species were collected at 2900–3000m, which is the highest locality known among the Zapotecanillus species. The collection site was located in a limestone area with sinkholes and karst topography, covered with a pine-oak forest. Soil temperature at the time of collection was 48°F (S.Peck, pers. comm.).

Externally, adults of Zapotecanillus montanus are similar to those of Zapotecanillus kavanaughi, Zapotecanillus iviei and Zapotecanillus pecki, but males of Zapotecanillus montanus may be distinguished from those of the other species by the shape of the median lobe (Fig. 48). See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in CMNH, point-mounted, labeled: \ MEXICO, Oaxaca, 3.5miles S of Suchixtepec, \ Ber.208, leaf litter, 3 June 1971 S.B. Peck collector\ CMNH\. PARATYPES (16 ex., 4♂2♀ were dissected), 6 ex. labeled same as a holotype; 6 ex. labeled: \ MEXICO, Oaxaca, 13 mi. N of Suchixtepec, 9500ft., ex. leaf litter, 4 June 1971, S. Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\; 4 ex. labeled: MEXICO, Oaxaca, 13.5 mi. S of Suchixtepec, 8000ft., ex. leaf litter, 3 June 1971, S. Peck\ THOMAS C. BARR COLLECTION 2011 Acc. No. 38014\ (deposited in CAS, CMNH).

The specific epithet is a Latinized eponym in the genitive case, and is based on the surname of Stewart B. Peck, Professor in the Biology Department of Carleton University, Ottawa, Canada, the collector of the type series of this species.

Mexico, Oaxaca, 3.5miles S of Suchixtepec.

Males of this new species are distinguished from those of other species of the genus by the shape of the median lobe (Fig. 51).

Size. Medium-sized for genus (SBL range 1.32-1.38 mm, mean 1.35±0.026 mm, n=5).

Habitus. Body form (Fig. 30) slightly convex, moderately elongate (WE/SBL 0.41±0.08), head of normal proportions for genus (WH/WPm 0.73±0.011), pronotum narrow compared to elytra (WPm/WE 0.72±0.009).

Color. Body monocolorous, rufotestaceous, appendages testaceous.

Microsculpture. Microlines partially effaced on disc of pronotum.

Prothorax. Pronotum (Fig. 35) relatively short (LP/LE 0.41±0.012) and slightly transverse (WPm/LP 1.24±0.030), with margins slightly sinuate and distinctly constricted posteriorly (WPm/WPp 1.36±0.043). Basal margin bisinuate near posterior angles. Contour of posterior angles slightly obtuse (108–118°) with 1-2 small denticles in front of the angles.

Elytra. Slightly convex, not depressed along suture, of moderate width (WE/LE 0.70±0.022). Margins subparallel at middle, slightly divergent in basal half, evenly rounded to apex in apical half, maximal width of elytra at midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally.

Male genitalia. Median lobe of aedeagus (Fig. 51), with elongate apex, rounded at tip. Dorsal plate 1 long, pointed apically and basally. Dorsal plate 2 joined to plate 1 at its apical third, where it forms a pronounced biapical protuberance. Ventral sclerite faintly sclerotized, barely visible. Right paramere rather long and moderately wide, with additional (3^rd) seta dorsally (Fig. 53). Left paramere without apical constriction (Fig. 52). Ring sclerite with short handle, widely rounded apically (Fig. 64).

Female genitalia. Spermatheca standard for genus.

The species is known only from the type locality in the Sierra Madre del Sur, in the surroundings of Suchixtepec (Figs 77 and 94, black flower).

Members of this species live at elevations of 8000-9500’ (2440-2900 m). At 8000’ (= 2440 m), beetles were collected in mixed pine-oak forest with Alnus, Carpinus, etc, and soil temperature at the time of collection was 56°F (S. B. Peck, pers. comm.).

Males of this species are easily distinguished from those of other members of the genus by the structure of the median lobe (Fig. 51) and setation of the right paramere (Fig. 53); and the geographical distribution of this species sets it apart from all its congeners. See also Fig. 90 for cladistic affinities.

HOLOTYPE, male, in KUNHM, point-mounted, labeled: \ MEXICO: Chiapas: Sierra Morena 16.16001°N, 93.60519°W, 1360m, 12-V-2008, ex. sifted leaf litter, 2° mesophil forest LLAMA08 Wa-A-01-1-all \ SM0833744 KUNHM-ENT \. PARATYPES (16 ex., 2♂3♀ were dissected), 9 ex. labeled same as a holotype, except barcode SM... numbers; 1 ex. labeled: \ MEXICO: Chiapas: Sierra Morena 16.15950°N, 93.60530°W, 1360m, 12-V-2008 sifted leaf litter, 2° mesophil forest LLAMA08 Wm-A-01-1 \ SM0839196 KUNHM-ENT \; 4 ex. labeled: \ MEXICO: Chiapas: Sierra Morena 16.15342°N, 93.60078°W, 1330m, 12-V-2008 ex. sifted leaf litter, 2° mesophil forest LLAMA08 Wa-A-01-2-all \ SM... KUNHM-ENT \; 2 ex. labeled: \ MEXICO: Chiapas, Mpio: Villa Corso, Ejido Sierra Morena, R. Biosfera La Sepultura, 16°09'10.6N, 93°35'25.1W, 1550m, 17–18.VII.2003, R. Anderson, mixed oak/pine forest litter, MEX1A03 110 \ SM... KUNHM-ENT \ (deposited in CAS, KUNHM).

The specific epithet is a Latinized eponym in the genitive case, and is based on the surname of John T. (Jack) Longino, Professor of the Biology Department of the University of Utah, and one of Co-PI’s of the LLAMA project, which provided the material on which the description of this species is based.

Mexico, Chiapas, Sierra Morena, 16.16001°N, 93.60519°W.

Adults of this new species can be distinguished from those of other species of the genus by the combination of small size and rufotestaceous color; and males can be further distinguished by the shape of the median lobe (Fig. 57).

Size. Small sized for genus (SBL range 1.01-1.12 mm, mean 1.08±0.038 mm, n=12).

Habitus. Body form (Fig. 31) slightly convex, slightly elongate (WE/SBL 0.40±0.10), head of normal proportions for genus (WH/WPm 0.76±0.021), pronotum moderately wide compared to elytra (WPm/WE 0.76±0.014).

Color. Body monocolorous, rufotestaceous, appendages testaceous.

Microsculpture. Partially effaced on disc of pronotum.

Prothorax. Pronotum relatively short (LP/LE 0.41±0.010) and moderately transverse (WPm/LP 1.28±0.035), with margins rectilinear and moderately constricted posteriorly (WPm/WPp 1.35±0.037). Basal margin oblique laterally. Contour of posterior angles obtuse (114–125°) with 0-1 small denticles in front of the angles.

Elytra. Slightly convex, not depressed along suture, of moderate width (WE/LE 0.69±0.015). Margins nearly subparallel, slightly divergent in basal forth, evenly rounded to apex in apical third, maximal width of elytra near midpoint.

Legs. 1^st male protarsomere markedly dilated apico-laterally.

Male genitalia. Median lobe of aedeagus (Fig. 57), with enlarged apex and neighboring part of ventral margin. Dorsal plate 1 small, pointed apically and narrowly rounded basally. Dorsal plate 2 located close to plate 1, in form of narrow stylet and shifted slightly apically. Ventral sclerites faintly sclerotized. Right paramere short and moderately wide (Fig. 59). Left paramere as in Fig. 58. Ring sclerite with short handle widely rounded apically (Fig. 67).

Female genitalia. Gonocoxite 2 comparatively short, with moderately curved blade and narrowly rounded apex (Fig. 71). Laterotergite with 7-8 setae. Spermatheca standard for genus (Fig. 76).

The species is known only from the type locality in the Sierra Madre de Chiapas (Figs 77 and 94, black triangles).

According to the label data, these beetles inhabit mesophyll and mixed oak/pine forests at low elevations.

In the structure of median lobe of males and geographical distribution, this species is only remotely related to its congeners. See also Fig. 90 for cladistic affinities.