Digonocryptus Viereck

Aguiar, Alexandre Pires & Ramos, Adriana C. B., 2011, 2846, Zootaxa 2846, pp. 1-98 : 6-12

Digonocryptus Viereck

Digonocryptus Viereck, 1913:373 . Type: ( Digonocryptus bidens Viereck ) = crassipes Brullé. Original designation.

Monogonocryptus Viereck, 1913:376 . Type: Monogonocryptus diversicolor Viereck. Original designation.

Odontocryptus Szépligeti, 1916:288 . Name preoccupied by Saussure, 1890 and by Cameron, 1903. Type: Odontocryptus variegatus Szépligeti. Monobasic. Description, keyed.

Digonocryptus: Townes and Townes, 1966:125 . Synonymy of Monogonocryptus .

Digonocryptus: Townes, 1970:330 Description, keyed, figure.

Digonocryptus: Yu and Horstmann, 1998:241 . Listed.

Digonocryptus: Kasparyan and Ruíz, 2005:170 . New species described from Mexico, keyed.

Diagnosis. All species show first tergite with spiracle near mid-length, fore tibia of female moderately to distinctly swollen, ovipositor quite straight, rigid, tip with lower valve partly enclosing upper valve, and the enclosing projection polished (in the similar Cestrus there is a scabrous area), fore wing cell 1+2Rs (areolet) closed, small, square or weakly rectangular, sometimes pentagonal; crossveins 2r-m and 3r-m are more or less parallel, receiving vein 2m-cu near or somewhat distad of middle; pleural carina of propodeum present between basal transverse carina and hind coxa. Other features include the clypeal margin medially with one or two small teeth; vein 1cu-a opposite vein 1M+Rs, or basad of vein 1M+Rs by not more than 0.35 its length; fore wing vein 2-Cu nearly as long as, or shorter than crossvein 2cu-a; hind wing vein 1-Cu distinctly longer than crossvein cu-a and the hind wing vein M+Cu is normally quite convex; T1 without a dorsolateral carina or with an incomplete one.

General description. FEMALE. The following features occur on females of all known species of Digonocryptus .

Head ( Figs 108–130). Mandible with two teeth with sharp mesal edges; ventral tooth equal in length or longer than the dorsal one, sometimes wider or more massive in size. A sulcus starting between the teeth, and extending at least to anterior 0.3 of mandible, but sometimes more, up to entire length of mandible, and extending medio- or dorso-longitudinally. Clypeus apical margin distinctly deflected, generating the apical area of the clypeus, which is usually black and often partially transversely strigate, medially always bearing two teeth or, more rarely, one tooth; ventral margin of deflection can be sharp or smooth and round. Species with two clypeal teeth show ventral margin of deflection medially usually weakly emarginated or incised, while somewhat swollen on each side, so that two weak lobes are normally distinguishable (as in Fig. 121); for species with one clypeal tooth the incision and lobes normally absent (as in Fig. 128). Malar space mat, microreticulate. Occipital carina always complete and always reaching hypostomal carina. A carina extending from anterior ocellus to base of torulus.

Mesosoma ( Figs 139–151). Epomia very short, reduced to a vertical ridge, equivalent to its anterior part. Medial part of dorso-posterior margin of pronotum with a distinct bell-shaped emargination, the margin of emargination thick and shinning, mesoscutum projected inside emargination. Notaulus always complete, though usually weak posteriorly. Central lobe of mesoscutum usually with longitudinal striation posteriorly. Speculum glabrous, polished, smooth. Posterior transverse carina of mesothoraxic venter strong and distinct laterally, but from short and weak to entirely absent ventrally. Anterior transverse carina of propodeum distinct and complete, medially bent or acuminated towards axillary through. Sculpturing on area in front of anterior transverse carina often smooth, or faint, or at least much weaker than general sculpturing of propodeum. Sternaulus reaching about 2/3 of length of mesopleuron; sometimes apparently complete (e.g., Digonocryptus chiriquensis ), but in fact meeting a different suture (not named), which runs from posterior transverse carina of mesosternum to mesopleural fovea. Hind tibia basal 0.2 narrowed, then slightly but perceptibly swollen, then progressively wider towards apex. Fore and mid femora with exact same color pattern, except dark colors generally slightly darker for mid femur, contrasting between different colors sometimes also a little more evident for mid femur. Sometimes, the long and stout setae on apex of hind tarsomeres 3–5, particularly those of t5, present what seems to be a spiral torsion, as a narwhal tusk.

Fore wing ( Figs 177–216). Venation largely uniform throughout the genus; 1M+Rs straight to slightly arched; 1m-cu convex, 2+3Rs concave; areolet about as tall as width of pterostigma, pentagonal, 2r-m and 3r-m parallel or converging anteriorly; a central bulla on 3r-m, 2+3Rs, and 2m-cu; 4Rs sinuous, longer than 2r; 2+3M straight to slightly concave, slightly longer than 4Rs, longer and parallel to 3Cu; 1-1A sinuous; 2-1A straight; 2Cua usually shorter, but sometimes as long as 2cu-a, which meets 2-1A at an angle of about 80º; 2m-cu straight. Posterior wing margin under vein 1-1A distinctly emarginated. Pilosity: basal cell with 3–5 rows near and along Sc+R, glabrous otherwise; 1 row along sub-basal cell, glabrous otherwise; distinct semicircular glabrous area on cell M1, near pterostigma, pilose otherwise (rarely entirely pilose, e.g., D. narratorius ), apical half of membranous area under vein 1A, and entire surface of all other cells. Variation: color patterns (from hyaline to uniformly infumate, or with a transverse stripe medially), 3Cu varying from as long as to twice as long as 2Cub (usually only slightly longer), 1M+Rs opposite or apical to 1cu-a.

Hind wing ( Figs 217–256). Venation highly uniform, with only slight variation throughout the genus. A bulla on posterior third of 1r-m; cu-a not connected to 1A; 1Rsb tubular and thick only at basal 0.15, otherwise nebulous, narrower; 2M, Cub and 2-1A nebulous. Pilosity pattern: Anterior wing margin with long ciliar hairs basally, changing suddenly to a line of small setae up to basal hamulus; with 3 (rarely 4–5) small setae beyond basal hamulus; membrane on Costal area mostly glabrous, except for pilosity on apical 0.2–0.5; usually 6–8 apical hamuli (number is proportional to wing size); all veins with regular line of pilosity anteriorly and posteriorly, often crossed in “X” over the vein; Sc+R with some stout setae on basal end; M+Cu with about 4 long, cylindrical, straight setae equally spaced; other setae on M+Cu, particularly the anterior ones on the apical half, somewhat flattened (less distinct in small species). All cells nearly entirely pilose, except sub-basal cell, mostly glabrous, with pilosity only in the apico-ventral angle, arranged in a more or less circular way; basal cell anteriorly usually with a distinct straight line of regularly spaced setae from base to apex (less distinct in large specimens), also with a few equally spaced stout and short setae among them. Vein 1Rsa touching but not continuous with Sc+R (noted under high magnification).

Metasoma ( Figs 91 –107). T1 usually polished, smooth; sometimes weakly microreticulate. T2 mostly and distinctly colliculate, matt, glabrous or nearly so; T3 as T2 but sculpturing weaker and somewhat mixed with alutaceous; T4–8 generally alutaceous, progressively weaker toward T8, which is partially or entirely smooth; T3–8 with fine pilosity, which is usually more abundant on T5 or adjacent tergites. T8 basally with a wide, shallow or deep depression, also somewhat folded down medio-longitudinally, in some cases generating a distinct crease ( Fig. 171).

Color. Species with clypeus or supraclypeal area yellow also have base of mandible and entire labrum yellow.

MALE. Generally similar to the respective females, but beginners may find them difficult to recognize because the most conspicuous features of the genus are absent in males, namely the dilated fore tibia, and the ovipositor features. In many species, the male gena is somewhat swollen ventrally, helping to characterize the genus. This is a subtle feature, particularly because of the reduced size of most males, but with practice it can be very helpful for the recognition of males of Digonocryptus . For species, most males can be recognized with the key to females provided in this work. Some secondary sexual differences, however, are intense, and somewhat characteristic. The most important are the following: total size considerably smaller than female size; antenna with many more flagellomeres than on female, the white stripe starting at a greater flagellomere number and usually covering more flagellomeres than on females of the same species. In many species, the gena is characteristically (for the genus) ventrally more swollen than on females, a feature originally noted by Bernardo F. Santos (UFES) (personal communication). Fore tibia simple, not inflated. Hind t5 simple, not deeply emarginated or bilobed, as on female; the apico-marginal line of 4–6 stout setae on each side absent. Propodeal apophyses and posterior transverse carina absent, or much less developed than on female. First metasomal segment long and slender when compared to the same segment in the female; also more straight, the downward bending observed between petiole and post-petiole on the female is attenuated or absent in males.

Species definition. The following characters are particularly useful in defining species of Digonocryptus , but are not listed in any particular order of importance, because their relevance will vary among different groups of species:

·Patterns of yellow on the lower metapleuron; this feature unites groups of species or, sometimes, might be characteristic for a given species (e.g., D. denticulatus ).

·Size and shape of the propodeal apophyses.

·Color pattern of all legs; it can be species-specific sometimes, as the unique pattern of D. chiriquensis (Cameron)

·The lateral shape of the first metasomal segment of females is often characteristic at the species level.

·Number of teeth on clypeal margin; it varies within the genus, but is quite constant intra-specifically.

·Yellow patterns on the propodeum, dorsally; this feature is unique and stable for several species, e.g., D. annulitarsis (Cameron) , D. chiriquensis , D. huntus sp. nov., D. meridensis sp. nov., D. tarsatus (Cresson) .

·Propodeal pilosity in species which have a black propodeum; this feature is characteristic in a group of otherwise very similar species, including those from the D. diversicolor complex.

The venation and color patterns of Digonocryptus wings do show some useful interspecific variation, but it is generally quite uniform, and therefore of limited taxonomic interest at the species level, as registered in Figs 177– 256.

The D. inflatus species complex. This complex is formed by Digonocryptus species with the mesosoma reddish and the metasoma dark with transverse yellow stripes, as in Figs 61, 63, 65, 67, 69, 137. It includes D. coloratus (Szépligeti) , D. inflatus (Brullé) , D. pulchripes (Cameron) , the synonymized D. rufithorax (see below), D. sutor sp. nov., and D. iageus sp. nov. Establishing a limit between these species can be very difficult for some specimens if this is performed only manually, that is, by subjective evaluation of their varying features. The cladistic analyses of all available female specimens ( Fig. 89), however, suggest that D. inflatus , D. pulchripes and D. iageus are indeed distinct species, whereas D. rufithorax is better treated as a junior synonym for D. inflatus , as proposed further below, in the respective description. The separation of D. coloratus and D. sutor is subtle (character:state 16:1 for D. coloratus ; also, D. coloratus has propodeum with eight complete strong transverse wrinkles from posterior transverse carina to base of petiole vs. nearly twice as much and much weaker wrinkles in D. sutor ). If only females are considered, the limits between these species are not very clear, suggesting they could be conspecific. However, the examined specimens do appear grouped on a particular clade ( Fig. 89), and, most importantly, the corresponding males for these taxa are considerably different, a fact which helps to support their isolation: males of D. sutor show metasoma with a wide white stripe on apical margin of T1, T3 and T7; males of D. coloratus have a narrow stripe on all tergites from T1 to T7.

Two male specimens, one from the Dominican Republic and one from the Virgin Islands (St. John), came from the AEIC grouped as a single species, together with males and females of what proved to be D. inflatus . Their distinctive color patterns suggest they might represent two undescribed species, but since males of Digonocryptus are usually less representative of the species than females, and since only one specimen is available for each presumed species, no further considerations will be provided in the present work for this material.

The following character coding scheme ( Table 1) was considered, corresponding to all informative external characters found to vary among the examined specimens. The respective character matrix is presented in Table 2. The resulting cladogram is shown in Fig. 89.

N° Description

01 Head main color: reddish [0]; black [1]

02 Orbital band from inconspicuous behind to interrupted between 11–1 to 11–3 h position [0]; complete or nearly complete, interrupted only about 11 h [1]

03 Supraclypeal area with yellow restricted to eye orbits [0]; supraclypeal area entirely yellow [1]

04 Ventral half of pronotum without yellow [0]; only with a fragmented yellow spot, covering at most 1/3 of the way from anterior center to posterior corner [1]; with a wide yellow stripe covering most of this area [2]

05 Mid tibia yellowish brown [0]; pale yellow to white dorsally, dark brown ventrally [1]; basal 0.3 white, otherwise dark brown, or entirely dark brown [2]

06 Hind tibia yellowish brown [0]; basal 0.2 white, otherwise dark brown [1]

07 T1 apex with a yellow stripe greatly narrowed medially, or interrupted, leaving one spot on each side [0], with a medial narrow yellow stripe, which does not reach sides [1]; with a wide, more or less uniform width yellow stripe [2]; narrow uniform stripe [3]

08 T2 and T3 apical margin with yellow stripe greatly narrowed to widely interrupted medially [0]; yellow stripe complete, not narrowed, sometimes widened medially, or present only on medial 1/3 [1]

09 T4–6 apical margin with yellow restricted to lateral sides, or greatly narrowed medially [0]; narrow yellow stripe present only on medial 0.7, not reaching sides [1]

10 Clypeal margin with 1 tooth [0]; 2 teeth [1]; teeth absent [2]

11 T2 behind thyridia without reddish spot [0]; with reddish spot [1]

12 Propodeal apophyses scale shaped [0], spine-shaped [1]

13 Mesopleuron unicolorous, reddish [0]; with yellow spot near wing base [1]

14 Mesosoma red or reddish brown [0]; yellowish brown [1]

15 Hind coxa without a pale yellow spot [0]; dorsally, centrally or basally with a pale yellow spot [1]

16 Posterior transverse carina medially faint or absent, marked mostly by weak scale-shaped apophyses [0]; complete and well developed [1]

The D. diversicolor species complex. Initially, this complex seemed to fit five closely similar but apparently well defined species: D. diversicolor , D. grenadensis , D. insularis , D. narratorius and D. arcaeus . Only with the cladistic analyses, however, it was possible to find clear support to propose two synonymies among them, D. grenadensis and D. insularis , both junior synonyms of D. narratorius . The females of the complex are characterized by a dense white pilosity covering the mesepimeron, carinal triangle, lower metapleuron, and propodeum ( Figs 79, 80, 83, 84), and often also the ventral half of mesopleuron; a complete orbital band, interrupted only at malar space ( Figs 79, 81, 83, 136); mesosoma main color black, with few small yellow areas (pronotum anterior margin and small spot on dorso-lateral margin, tegula, subalar prominence, scutellum and post-scutellum) ( Figs 133, 135, 136, 152, 153), and metasoma mostly or entirely orange ( Figs 133, 135, 136, 152, 153). Males can be recognized by a similar set of features, but the metasomal color pattern is more variable (e.g., Figs 91, 92, 93).

outgroup species.

Structurally, these species seem to show consistent differences among them, but they also display a rich and intricate set of individual variations of their basic color pattern, rendering species interpretation even more difficult and subjective. In order to objectively assess these variations, the cladistic analysis was very helpful, allowing the consideration of all available specimens of all species of the complex, while also taking into consideration all observed morphological variation among them.

The results ( Fig. 90) suggest an interpretation of D. diversicolor as a clade of specimens which is isolated from the D. arcaeus clade. In the strict consensus tree, one specimen [D28] seems somewhat intermediate between D. arcaeus and D. diversicolor ( Fig. 90). However, it is isolated from all D. diversicolor specimens by lacking three synapomorphies of that species, namely character-states 2:2 (supraclypeal area without a medio-longitudinal stripe or, if present dorsally, very short, not reaching parantennal impression), 12:0 (propodeal pilosity behind posterior transverse carina uniform), and 19:1 (T2 apical margin with transverse yellow stripe). The same tree shows a distinct clade for D. narratorius , with D. grenadensis and D. insularis grouped together, and therefore clearly suggesting the synonym.

The following character coding scheme ( Table 3) corresponds to all observed informative variation of external characters in the studied specimens. The respective character matrix is presented in Table 4.

outgroup species.