Journal of Science: Vol. 10 | Article 111 Serna and Mackay

A descriptive morphology of the Procryptocerus (: Formicidae)

F. Serna1,2a, W. Mackay1,2b

1Grupo Sistemática de Insectos Agronomía (SIA). Línea Taxonomía de insectos. Universidad Nacional de Colombia. Facultad de Agronomía. Museo Entomológico Universidad Nacional Agronomía Bogotá (UNAB). Cra. 30 # 45-03. Bogotá, Colombia Downloaded from 2Department of Biological Sciences. The University of Texas at El Paso. El Paso, TX, USA. 79968

Abstract Morphology is the most direct approach biologists have to recognize uniqueness of insect species as compared to close relatives. of the genus Procryptocerus possess important morphologic http://jinsectscience.oxfordjournals.org/ characters yet have not been explored for use in a taxonomic revision. The genus is characterized by the protrusion of the clypeus forming a broad nasus and antennal scrobes over the eyes. The toruli are located right posterior to the flanks of the nasus opposite to each other. The vertex is deflexed posteriorly in most species. An in-group comparison of the external morphology is presented focusing on the workers. A general morphology for gynes and males is also presented. Previously mentioned characters as well as new ones are presented, and their character states in different species are clarified. For the metasoma a new system of ant metasomal somite

nomenclature is presented that is applicable to Aculeata in general. Finally, a Glossary of by guest on September 29, 2015 morphological terms is offered for the genus. Most of the terminology can be used in other members of the Formicidae and Aculeata.

Key words: Hymenoptera, Aculeata, Formicidae, Cephalotini, qualitative characters, external anatomy, Correspondence: a [email protected], b [email protected] Received: 1 February 2009, Accepted: 1 October 2009 Copyright : This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 10, Number 111 Cite this paper as: Serna F, Mackay W. 2010. A descriptive morphology of the ant genus Procryptocerus (Hymenoptera: Formicidae). Journal of Insect Science 10:111 available online: insectscience.org/10.111

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Journal of Insect Science: Vol. 10 | Article 111 Serna and Mackay

Introduction a template for a revisionary study of the entire group following the recommendations The genera Procryptocerus and Cephalotes suggested by Bolton (2007): “… [not] make comprise the tribe Cephalotini (de Andrade an unwarranted assumption that previous and Baroni-Urbani 1999. Emery (1922) authors have already investigated all the demonstrated that the tribe possesses the useful characters...”; “Initial dependence on synapomorphic anatomical trait of mushroom- previous publications has a strong tendency to head-shaped proventricular valves (Kempf restrict the scope of a new investigation…”; 1951) (Figure 1). This observation is “… develop a personal insight into [the] supported by the studies of de Andrade and morphology and variation that is not unduly Baroni Urbani (1999). Procryptocerus was influenced by what has previously been created by Emery (1887) to include species of published.” Knowledge of morphology and Neotropical ants that were considered similar anatomy is incomplete for all species. New Downloaded from to those of the Paleotropical genus Cataulacus characters must be discovered, and old of the tribe Catalaucini (Kempf 1951). characters tested. Morphological descriptions Procryptocerus is a lineage composed of are thus essential components of our about 80 species inhabiting rainforests from understanding of species and their diversifi- http://jinsectscience.oxfordjournals.org/ the Isthmus of Tehuantepec in Mexico to cation (Wheeler 2008, Bert Hölldobler, northern Argentina. Due to their cryptic Universität Würzburg, personal communi- habits, living inside twigs, these ants are cation). rarely collected (Mackay and Vinson 1989). At present, most species are known from This approach is divided into two sections. In Central America, Colombia and Brazil. the first part, a diagnosis is presented that expands previous morphological diagnoses of

Procryptocerus has been the object of two the genus provided by Kempf (1951) and by guest on September 29, 2015 revisionary studies. Kempf (1951) revised the Longino and Snelling (2002). Additional entire genus and Longino and Snelling (2002) observations, current morphological termi- the Central American species. Kempf (1951) nology, and figures are part of this new recognized 28 species, and 8 subspecies, diagnosis. A unification of the terminology is while for Central America Longino and proposed within ants and with other Aculeata Snelling (2002) recognized 14 species, regarding the specialized system used to name described four new species, synonymized two the metasomal somites (abdominal (Abd) species, and elevated two subspecies to somites II to pygidium) (see Materials and species level. Currently, 56 nominal taxa are Methods, including definition of cinctus). included in the genus (Bolton et al. 2006). In the second part, a selected Glossary is Procryptocerus ants possess notoriously offered containing terminology appropriate variable morphology. Different characters, for Procryptocerus. The provided terminology such as propodeal spine length, form of the might also be used for descriptions as well as clypeus, type of sculpture, and other such identification keys of other taxa of ants. characters vary remarkably, sometimes even Working on the study of morphology and the within the same species. A diagnostic mor- associated terminology is a constant necessity phology of the genus is proposed to be used as in order to unify criteria for the basic

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descriptive work in taxonomy and 2006), Triplehorn and Johnson (2005), Wilson comparative biology. (2003) and Ward (1999, 2004).

Materials and Methods Terminology Terminology indicated within parentheses and Worker, gyne and male specimens of quotation marks, e.g. (“girdling constriction”) Procryptocerus scabriusculus from CWEM has been avoided. Words within brackets and (William and Emma Mackay Collection, El italized, e.g. [mayri] are examples of species Paso, Texas, USA) were drawn at 60X power possessing the specific character state pointed with a Wild Heerbrugg microscope using a out; accordingly, this bracket use does not grid and a micrometer. Some structures were indicate that the defined character state is cleared in potassium hydroxide (10%) for 36 restricted to the examples given. No examples hours. To analyze the metasomal sclerites, the are given when the character state explained is method in Bolton (1994) was followed. To fairly common. Numbers within brackets Downloaded from show differences in sculpturing, structures of following the capital letters FS_, [FS_01], different species were also drawn. A diagnosis [FS_02], [FS_03], etc., indicate species level of males is modified from Kempf (1951). taxa in the process of being described. When Since few, and in some cases are not available hyphenated sculptural states are used, i.e. http://jinsectscience.oxfordjournals.org/ for study males in the genus, this “foveoate-costate”, both forms of sculpture morphological diagnosis emphasizes females are present as intraspecific variation. In the and is concentrated in qualitative characters. Discussion sculptural terminology is orga- Terminology used for positions and nized as interpreting the form of sculpturing orientations is explained below. For the study from the smallest to the largest. of exoskeletal morphology, the main literature resources were Snodgrass (1935), Bohart and Specific positions such as basal, proximal,

Menke (1976), Gauld and Bolton (1988), distal, apical, apicolateral, apicomesial are by guest on September 29, 2015 Bolton (1994), and Ward and Downie (2005). used exclusively for appendages such as Sculptural terminology is from Sparks (1941), buccal appendages, antennae, wings, legs, or Harris (1979), Torre-Bueno (1989), Brown genital appendages (Figure 2). Indications of (1979) and Hölldobler and Wilson (1990). positions such as “propodeal base”, “gastral Vestitural terminology is from Sparks (1941), base”, “base of declivity” are avoided since Hölldobler and Wilson (1990), and Ward they are referring to structures on the mesion, (2004). Although Procryptocerus ants are and not to appendages. Other specific mostly black, variation in color is included to positions (anterior, posterior, dorsal, ventral, help distinguish some forms. Specific lateral, mesial, etc.) are used for body parts; terminology is selected from different relative positions (with the adverbial ending publications used for descriptions of ants and ad meaning toward, such as basad, distad, other Apocritans: Snodgrass (1935), Gauld anteriad, cephalad, posteriad, caudad, laterad, and Bolton (1988), Nichols (1989), Gordh and mesiad, anterodorsad, anteroventrad, Headrick (2000), Hölldobler and Wilson posterodorsad, posteroventrad, etc.), direc- (1990), Mackay (1991, 1993), Bolton (1990, tionality (mesially, laterally), and extended 1994, 2003), Agosti et al. (2000), Longino and positions, that involve two or more regions Snelling (2002), Mackay and Mackay (2003, (dorsolaterally, dorsosternally, dorsoventrally, lateroventrally, anteroposteriorly, posteroan-

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teriorly, lateromesially, etc.), are used for both laterally. The lateropropodeum is the lateral the mesion and the appendages. In relative area (laterotergite) of the anteropropodeum (ad) positions, such as lateroventrad, the containing the propodeal spiracle. The prefix “latero” emphasizes that the structure posteropropodeum is located beneath the or character state is lateral and the suffix propodeal spines; it is the posterior vertical or “ventrad” indicates that it is found in the declivitous area of the propodeum. direction of venter. The opposite applies to ventrolaterad or other combinations of Metasoma (Figures 37, 38, Table 1). prefixes and suffixes indicating relative In referring to metasomal somites in ants, positions. usually two different systems are super- imposed (Bolton 1994). A general system In the sense used here, ante means before regards homologous abdominal somites (Abd) (anterior to) the referred structure (e.g. throughout the Hexapoda. In the Formicidae, antepropodeal refers to a structure anterior to as is it in the entire Apocrita, Abd I is part of Downloaded from the propodeum), and antero refers to the the second tagma or the mesosoma, which is anterior portion of the actual structure. formed from the thorax plus Abd I tergite. The Anteropropodeum refers to the anterior region remaining abdominal somites form the third on propodeum. Nevertheless, very common tagma starting at Abd II (petiole). A second, http://jinsectscience.oxfordjournals.org/ literature uses of the prefix “pre” are not specialized (functional) system divides the changed when, for example, making reference metasomal somites into a petiole, postpetiole to presclerites as presternite, pretergite, etc. and gastral segments (Bolton 1994, 2003). Because different groups of ants contain The use of terminology for shapes is quite forms with one- or two-petiolate metasoma, useful for describing different structures. A the current specialized system of metasoma combination of technical and common (not nomenclature uses the name “gaster”

universal) names describing shapes is present inconsistently and incongruently with homo- by guest on September 29, 2015 in the literature. For instance, terms such as logous somites in the non-formicid crescentiform, fusiform, disciform, etc. are Apocritans. This situation shows that the technical and therefore universal. Terms such development of a consistent system of naming as “neck” for a part of the antennae, “cheeks”, specialized metasomata has passed behind the “apron”, etc. are not technical, not universal terminology for prosoma or mesosoma. and therefore are avoided. Bolton (1990) introduced the term helcium. A second helcium is characteristic of two- Propodeum (Figures 9, 10, 15, 37, 38) petiolate metasomata; when it is not present, The propodeum is the first abdominal tergite no specialized term is available for Abd III, fused to the thorax, which together comprise and hence the term postpetiole is the mesosoma. The propodeum is differen- inconsistently used between different castes tiated into the anteropropodeum and the and subfamilies. Occasionally, authors have posteropropodeum. In turn, the anteropro- had to explain the need of petiole and podeum is divided into the dorsopropodeum postpetiole in males without helcial sclerites and the lateropropodeum. The (see de Andrade and Baroni-Urbani 2003). A dorsopropodeum is the dorsal area of simple solution to the inconsistent use of the propodeum, anterior to propodeal spines and terminology regarding postpetiole and gastral containing the anteropropodeal processes somites in ants would be to abandon the

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specialized system of metasoma vocabulary. The metasoma comprises two general However, the use of a specialized system has subdivisions: the anterior metasoma shown interesting advantages in the constituting the petiole (Abd II), and the comparative morphology of ants (see Bolton posterior metasoma composed of metasomal 2003), and is applied in recent classifications (mtm) 2 (Abd III) to pygidium. In the (Bolton 2003, Perrault 2004, Ward 2007). posterior metasoma, Abd III-IV (mtm 4-5) Therefore, improvements to the specialized constitutes the middle metasoma, Abd V to metasomal terminology are desirable. This pygidium constitutes the caudal metasoma. work proposes a proposal of reconciliation The petiole (mtm 1) contains the cinctus 1 into a single morphological specialized system (cinctus: constriction between pre and for what we believe are homologous postsclerites, see below), the mtm 2 contains metasomata within ants and other Aculeata the cinctus 2, and the mtm 3 may or may not (Table 1). contain the cinctus 3. The petiole (Abd II) and mtm 2 (Abd III) are joined by the first Downloaded from In Apocrita, the metasoma is composed of helcium (Bolton 1990) in all ants. In the Abd II to the caudal segment, or periproct middle metasoma, the mtm 2 and 3 (Abd III- (Snodgrass 1935). Literally, “gaster” means IV) may or may not be joined by a second stomach (Brown 1979). However, the word helcium. The mtm 2 is usually considered the http://jinsectscience.oxfordjournals.org/ “gaster” has widely been used in the literature postpetiole when separated from mtm 3 by the of Apocrita referring to the external third helcium-cintus 3 complex. This state is seen tagma minus the peduncle (“petiole” in no in females of Myrmicinae, ants) (Bohart and Stange 1965, Naumann Pseudomyrmecinae, and others. In other 1991). The gaster constitutes the abdomen considerations of a postpetiole, the mtm 2 is without Abd I (propodeum) and the peduncle separated from mtm 3 by the cinctus 3 only of Abd II (Naumann 1991). (e.g. see de Andrade and Baroni-Urbani 2003

for Proceratium). This state is known to occur by guest on September 29, 2015 The Abd II (petiole) is always specialized in in males of Myrmicinae, Pseudomyrmecinae, ants (Bolton 1994). The petiole is also found and in both sexes of Ecitoninae (including in some groups of Tiphiidae, Scoliidae, Cheliomyrmicini), Ectatomminae, Vespidae, Mutillidae and other Aculeata, Amblyoponinae, Cerapachyinae, , where Abd II is a specialized somite as well. Paraponera clavata, Proceratium, and others. The petiole may be nodiform, squamiform or a much reduced subcylindrical segment For the posterior metasoma, we propose the (Bolton 1994). Pedunculate, sessile and use of the name gaster when a postpetiole is subssesile petioles in ants are, in general, not present (e.g. Formicinae, Dolichoderinae), artifacts of the anteroposterior displacement of and the use of postpetiole and opisthogaster the petiolar nodus. In groups with a when at least a cinctus 3 is present. The pedunculate petiole (e. g. Pheidole or posterior metasoma is a gaster when its Solenopsis), the node is posterior. In sessile metasomata 3 to pygidium (Abd IV to petioles (such as in Procryptocerus), the pygidium) are combined as a unit (e.g. tergum is augmented anteriorly (nodal) Formicinae, Dolichoderinae); i.e. the gaster usually forming an anterior nodal truncation. constitutes the metasoma minus the petiole when a postpetiole is absent. The opisthogaster constitutes the combined caudal

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metasomata 4 to pygidium (Abd V to posterior to the second helcium (sensu Bolton pygidium); i.e. the opisthogaster is the 1990), in two-petiolate ants. When the cinctus posterior metasoma minus postpetiole. Even 3 is present in some of the sensu stricto single in few cases, when it is not clear to associate petiolate ants, it is a very fine sulcus (e.g. gaster and opisthogaster to general different poneroids groups), or almost absent subfamilies, tribes or genera, the two terms (e.g. Odontomachus). Few genera such as would stay associated to the already explained Leptanilliodes possess more than three cincti. metasomata.

In summary, we propose a consistent Results specialized (functional) and morphological system for naming homologous metasomata in Workers ants. When the cinctus 3 is absent in the metasomite 3 (Abd IV), we propose using Diagnosis Downloaded from gaster for the combined metasomata 3 to Frontal carinae posteriorly divergent; malar pygidium. When a postpetiole is considered, space not covered dorsally by frontal carinae; due to the presence of at least the cinctus 3, clypeus protruded developed into a broad the posterior metasoma is divided into nasus lying between toruli; antennal scrobes http://jinsectscience.oxfordjournals.org/ postpetiole and opisthogaster. impressed laterally above eyes, limited by frontal carinae dorsally (Figures 3, 4), Cinctus 1, 2, 3 (pl. cincti) (Figures 3, 9, 16). extending from nasus to vertex; vertex Cincti are the anteriad sulci, often in the shape deflexed (truncate) posteriorly in most of a belt or girdling constriction, located on species; toruli located right behind nasal metasomal 1 (Abd II = petiole), metasomal 2 flanks opposite to each other; Abd V (Abd III = postpetiole) and metasomal 3 (Abd postergite visible from above. Adult coloration mostly black, body variously

IV) somites. The sulci separate the pre and by guest on September 29, 2015 post sclerites. Cinctus 1 is usually a dorsal, sculptured; workers monomorphic, ranging very slender sulcus concealed between the from 3.5-8.5 mm (Kempf 1951), gynes from posteropropodeal lobes, anterior to the 3.7 mm [schmitti] to 9.5 [mayri part]. peduncle and sternopetiolar process in petioles possessing these structures. In some groups Head (Figures 3, 4, 5, 8, 9) (e.g. Pogonomyrmex) there is no apparent Prognathous, usually in anteroventrad position cinctus 1. In such cases, cinctus 1 is in preserved specimens (Figure 9); interpreted as the anteriormost portion of the prognathism formed by combination of hypostomal, genal and postgenal bridges. In peduncle located between the posteropropodeal lobes. In general, in sessile full frons (“full face”) view, subtrapezoidal, broadened posteriorly (Figure 4), circular metasomata (e.g. Procryptocerus) cinctus 1 is located directly anterior to the spiracle. The [nalini] (Figure 23), subcircular [regularis, cinctus 2 is located in metasomal 2, anterior to pictipes] or subquadrate [belti (Figure 20), FS_02]; in lateral view, more often elliptic; the ventropostpetiolar (“subpostpetiolar”) process in metasomal 2 that possesses such mandibles, anteclypeal carina and clypeus process. The cinctus 3 (the “girdling anteriorly positioned; frontoclypeus (“face”) constriction”- Bolton 1994) is located between anterodorsally extended, comprising epistoma the pre and postsclerites of the metasomal 3, (anteclypeus, plus clypeus) and frons; frons dorsal; vertex posterior, opposite to nasus,

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often orthogonal to frons (deflexed); in dorsal facial fovea (malar depression), when present (full frons) view, frontoclypeal (epistomal) [mayri, batesi, carbonarius, rudis] (Figure 24) suture and frontovertexal margin located in encircling part of clypeogenal suture, and nearly same plane; malar space (Gauld and often part of antennal fovea (Torre-Bueno Bolton 1988) extended dorsolaterally, laterad 1989); malar tumulus (Figure 24) anteriad to to antennal scrobe and anteriad to eye (Figures eye and limiting facial fovea laterally [batesi, 3, 4, 9); hypostomal bridge and genal bridge rudis]; frons anteriorly delimited by faint ventral (Figure 7), opposite to frons, occiput frontoclypeal (epistomal) suture or faint and occipital foramen posteroventrad, frontal triangle (Figure 4), laterally by frontal postgenal bridge ventroposteriad, posterior to carinae, posteriorly by frontovertexal margin; genal bridge, with the two areas meeting in a frontal carinae diverging from clypeus to curve or an angle behind eye; anteclypeal vertex, straight, uni or bilobate [mayri, batesi] carina (Figure 4) indistinct, emarginate, or (dorsal view) into frontal lobe and frontal bilobate; clypeus differentiated into discal carina posterior lobe(postfrons lobe), posterior Downloaded from (central) and truncate strikingly orthogonal lobe deflexed posteriad in some species lateral nasal flanks toward toruli (Figures 3, 4, forming an angulate scrobe [rudis] (Figure 9) [eladio, pictipes, goeldii, marginatus], or 24); frontal lobe indistinct (not flanged) trapezoidal, not orthogonally flanked, being (Figures 3, 4, 9, 17-19, 23, 29) [scabriusculus, http://jinsectscience.oxfordjournals.org/ wider on lower lateral area toward malar hylaeus, brazilian species] or distinct [rudis, space and forming invaginated antennal fovea mayri, batesi] (Figures 20-22, 24); antennal (Figure 24) [batesi]; frontoclypeal (epistomal) scrobe lateral, under frontal carina, not visible suture obsolete between toruli, turning down in full frons view, formed of long, deep, wide laterad between nasal flank (located lateromesial invagination (groove), extended anteriorly) and torulus (located posteriorly); anteroposteriorly, starting at lateral fovea, epistomal suture lateral between clypeus and passing over eye, and terminating in notch on

torulus, not grooved [paleatus], faintly vertex at frontovertexal corner level (Figures by guest on September 29, 2015 grooved [mayri], or variably grooved and 9, 24), dorsally delimited by frontal carina and forming clypeo-torular sulcus [carbonarius, ventrally by dorsal ocular suture or anteocular some rudis] (Figures 9, 24), continuing costula; eye lateral, infra antennal scrobe downward to pleurostoma, anteriad to malar (head in profile), slightly posteriad, or space, sometimes indistinct and forming intermediate between torulus and vertexal vanishing clypeomalar (clypeogenal) suture margin; eye globular [convergens, rudis], flat (Figures 3, 4) separating narrow premalar [FS_03], rounded not-protruded [marginatus], space from malar space; anterior tentorial pit ellipsoid [scabriusculus, kempfi], dorsally located into clypeomalar suture, anterior to depressed [adlerzi, sampaioi]; vertex lateral fovea (Gordh and Headrick 2000); deflexed, often delimited anteriorly from frons nasal flanks anterior to toruli; discal clypeus by transverse frontovertexal margin, protruded into broad, nasus (Figures 3, 4, 9) in posteriorly by occipital carina, and laterally by approaching frons; nasus describing in profile posterior notches of antennal scrobes [batesi, short anterodorsal curve at toruli level; lateral rudis, mayri, virgatus]; frontovertexal margin fovea beneath torulus (not visible in dorsal distinct throughout forming fastigium (Figures view) and posterior to clypeomalar suture, 3, 4, 9, 14, 15, 16-19, 25, 27, 29) [impressus, marking antennal scrobe at most anterior point paleatus, regularis, adlerzi, scabriusculus], and receiving lamella of scape shaft base; indistinct throughout [coriarius, schmitti]

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(Figure 23), medially indistinct and laterally victoris, seabrai, FS_03] or relatively distinct [mayri, clathratus] (Figures 20-22), or concealed by torulus [carbonarius]; scape frontovertexal margin outline (frontal or stalk truncate basally [attenuatus], often posterior view) straight [eladio, kempfi, proximally terete (narrow), wide [attenuatus, subpilosus], slightly medially notched nalini], or slightly tapered [scabriusculus, [scabriusculus in part, convergens] (Figures 4, convergens], basally uni or bilamellate (broad, 19), crenate [clathratus, impressus, thin, flanged carina) overlapping condylar marginatus, spiniperdus] (Figure 18), constriction [rudis], or ecarinate [schmitti]; crenulate [paleatus], convex [coriarius, scape lateral axis (lateral stalk) not completely nalini] (Figure 23), slightly convex-crenulate covered by frontal carina when [paleatus], biconvex [adlerzi] or flanged- accommodated into antennal scrobe; scape biconvex [adlerzi] (Figure 27); frontovertexal shorter than scrobe length, terminating corner with (Figures 3, 4, 9, 16, 17, 20, 22, between eye and vertex; funiculus 10- 27) or without [coriarius,nalini] (Figure 18) segmented, second funicular segment shorter Downloaded from angulate processes; vertex concave than first or third, three distal segments [clathratus, marginatus, paleatus, impressus, compose club, apical segment subconical. spiniperdus], flat [scabriusculus, lepidus, balzani] or slightly convex [coriarius]; Mesosoma (Figures 3, 9-11, 26-30) http://jinsectscience.oxfordjournals.org/ hypostomal tooth lobose. Mandible (Figures Subcubic, trunk-shaped, dorsally convex or 5-7) subrectangular, possessing scrobe flat. Tergal, pleural (meso and meta) and ventrally and trulleum dorsally, massive, propodeal sclerites fused into notosternal- medially turned; inferior margin (lateral view, propodeal capsule. Pleura subparallel or mandible closed) ventrad [generally species at mesially inflected between meso and low elevations], anteroventrad [most Andean metapleura. Pronotum hood-shaped, covering species over 600 m]; apical (“dentical”) more than 1/3 of dorsal and lateral areas of

margin medially directed, possesing major mesosoma, forming dorsopronotum and by guest on September 29, 2015 distinct, infra, apicolateral (“apical”) tooth; lateropronotum (= pronotal side panel - Gauld preapicolateral tooth (tooth number 2) half and Bolton 1988) (Figure 9); dorsal profile size of apicolateral; tooth number 3 [obsolete from markedly convex [gibbossus] to in pictipes, mayri] half size of number 2 horizontal (dorsally flat) [paleatus, [FS_01]; tooth 4 obsolete or absent; supra, impressus]; dorsopronotum and apicomesial (“basal”) angle acute. Maxillar- lateropronotum divided by pronotum labial palp formulae 4-3 [FS_01, carbonarius] dorsolateral margin (Figure 9); (Figure 8), 6-3 (Kempf 1951); palpifer almost lateropronotum subtriangular with vertex at half size of first palpomere; first and second level of procoxal base, sometimes forming palpomeres same size, apical (fourth) longest. inferior lamella (inferior pronotal process - Antenna 11-segmented; scape comprises Figure 10) flanking procoxa basally [Andean short, basal radicle and long, distad shaft; species], both structures (inferior pronotal radicle divided into condylar bulb (Bolton process and procoxa base) probably forming 1994) (Figures 4, 14) (inserted into torulus) stridilum; lateropronotum overlapping and condylar constriction (“neck”), which propleuron in most species (Figure 9) and functionally fits into posterior notch of torulus forming pronotal lobe posteriorly (Figure 9), during anteroposterior movement of scape; lateropronotum mesially inflected conforming condylar bulb anteriorly visible [attenuatus, humeral angle anteriad (Figure 3) and

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humerus (elongate, narrow anterior area Andean species], truncate, square [adlerzi, connecting with neck dorsally); humeral regularis], or falcate [victoris] epicnemial inflection forming lateral carina limiting process (Figures 10, 12, 30) flanking procoxae humerus and lateropronotum (wide lateral externally, both structures probably forming panel); lateropronotum flat discally (Figure 9) stridulating organ; notopropodeal fusion often or slightly inflected (Figures 29, 30) receiving marked by groove and lateral excavations disciform profemur [impressus], both [spiniperdus, eladio, adlerzi] (Figure 3) or structures probably forming stridulating lateral excavations only [sampaioi]; organ; ventro-propleurites separated by ventro mesopleuron usually inflected (mesosoma medial suture (Figure 11), sometimes constricted) receiving femora downward apparently fused; ventropropleurite and notopropodeal excavation; metapleural gland humerus forming protruded or flat area scrobe (Figures 10, 12) superior to metacoxa, coupling postgenal bridge when head deflexed extending posteroanteriorly from metapleural downwards; prosternum between procoxae, gland bulla (Figures 10, 12) to mesopleural- Downloaded from posteriad to ventropropleurites; mesonotum coxal excavation (mesopleural coxal process - flat [pictipes] or convex [sampaioi], laterally Snodgrass 1935- inflected into excavation) fused to anepisternum (Figures 9, 10), (Figure 10); metapleural gland scrobe anteriorly delimited by vestigial promesonotal canalicular, channel delimited superiorly and http://jinsectscience.oxfordjournals.org/ suture (Figures 3, 9, 26, 27), extending inferiorly by two longitudinal carinae, inferior posteriad until meeting notopropodeal fusion carinula flanking metapleural gland slit (see Discussion), usually possessing lobose or ventrally; metapleural gland extending spiniform lateral process (mesonotal process) dorsoposteriorly, turning downward (Figures 3, 9, 27); promesonotum possessing ventroanteriorly forming metapleural gland lateral excavations between pronotal lobe and slit (Figure 12); slit very narrow, bicarinulate, mesonotal process; lateropronotum and running ventrally posteroanteriorly from

mesopleuron separated by open, narrow, metapleural bulla to mesopleural-coxal by guest on September 29, 2015 nearly straight or sinuate lateropronotal- excavation; propodeum (first abdominal (Abd mesopleural suture continuing dorsally, I) tergite) divided into anteropropodeum and forming promesonotal excavation between posteropropodeum, first subdivided into pronotal lobe and mesonotal process, dorsal (dorsopropodeum) and lateral connected to promesonotal suture (Figure 3); (lateropropodeum) areas (Figures 9, 10); anapleural sulcus (“anterior oblique sulcus”) lateropropodeum including spiracle, (Figure 19) down promesonotal excavation, posteropropodeum (declivitous face) (Figures dividing smaller anepisternum supra and 9, 10) under propodeal spines terminating at larger katepisternum infra (Figures 9, 10); posteropropodeal lobes (propodeal lobes) katepisternum extended lateroventrally, (Figure 12); dorsopropodeum usually ventrally forming epicnemium (Gauld and horizontal in profile [FS_01, scabriusculus] to Bolton 1988) separating pro and mesocoxae; convex [hylaeus, sampaioi], same plane as epicnemial carina (“omalus” sensu Bohart and mesonotum [pictipes, lenkoi, sampaioi] or Menke 1976) (Figures 10, 12) and lower than mesonotum [clathratus], expanded epicnemium forming shallow-concave surface anterolaterally into somewhat anteropropodeal receiving procoxa posterior face; epicnemial process (Figure 3); propodeal spines (Figure carina projected anteriorly into distinct 3) horizontal and parallel [kempfi], divergent laminate, variable shaped, subcircular [most and upturned [rudis], parallel upturned

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[clathratus], parallel upturned-curvate pectinate probasitarsus; meso and [eladio], parallel down-turned [scabriusculus metabasitarsi cylindrical and longer; postarsus part] (Figure 9), or different sizes within same (“pretarsus”) formed by bilobed padded population [scabriusculus part]; propodeal arolium and bifurcate curvate claw. spiracle tubulose, downward on lateropropodeum between anteropropodeal Metasoma (Figures 3, 9, 12, 13, 15, 16, 26- process and propodeal spine base, commonly 34, and Table 1) directed posteriorly, often accommodated into (Justification of terminology used here for lateropropodeal excavation, internal margin third tagma in Metasoma can be found in generally fused to excavation, lateral margin Materials and Methods and the Glossary. usually free; posteropropodeum sometimes Petiole (metasomal 1 = Abd II), first forming continuously concave descending specialized metasomite articulated to declivity until reaching posteropropodeal propodeum by manubrium (Perrault 2004), lobes, somewhat vertical or slightly diagonal composed of tergite and sternal presclerite Downloaded from supra, shorter infra (between posteropropodeal (Figure 12), forming syntergosternite (tergite lobes), supra and infra areas rarely same and sternite fused), sessile, constricted length [scabriusculus part] (Figure 9); anteriad into cinctus 1 (cinctus: constriction posteropropodeal lobe posterior to bulla between pre and postsclerites, see under http://jinsectscience.oxfordjournals.org/ (Figure 12); metacoxal cavity ental (Figure Metasoma in Materials and Methods) (Figure 12); legs similar to male’s (Figure 47), 12), nodiform, narrower than distance procoxa trunk-like augmented basally, twice between propodeal spine bases, subcylindrical size of meso or metacoxa; profemur tectiform [kempfi], slightly wider anteriad, or barrel- (roof-like), securiform (triangular) in cross shape [eladio, batesi], usually without dorsal section (clear vertices on inferior side), or lateral excrescences or projections; node equilateral, or ventral side narrower than anterior face reduced [nalini] or more

anterior and posterior sides [belti, eladio], commonly truncate forming nodal truncation by guest on September 29, 2015 fusiform (spindle-shaped) [mayri, batesi], or (Figures 3, 32) opposite to and functionally compressed disciform (disc-shape) received by posteropropodeum, often [impressus] with dorsal margin carinate delimited by nodal dorsolateral margen; nodal (keeled) [impressus, paleatus] and concave truncation convex, straight, curvate anteromesially (entad) and proximally supraposteriad [adlerzi], concave [coriarus, [impressus, paleatus], or convex; slightly sampaioi], or absent [nalini], petiolar summit convex posterolaterally; meso and anteriad (Figures 3, 29, 31), midway [hylaeus] metafemora commonly tectiform, ventral side or posteriad [seabrai]; sternopetiolar weakly concave, separated from trochanter by (“subpetiolar”) process between cinctus 1 and small, dorsad, cuneiform prefemur (Kukalová- node (Figures 29, 31), and usually laminar- Peck 1991) (“trochantellus”) (Figures 9, 47), lobose [scabriusculus, rudis, mayri] or (male profemur elongate fusiform (Figure obsolete, petiole posterior foramen margin 47)); tibia subcylindric, possessing four (lateral view) sinuate (Figures 9, 10) or poorly defined panels, anteriad and posteriad vertically set off, spiracle anteroventrad. wider than ventromesiad (flexor) and Postpetiole, second specialized metasomite dorsolaterad (extensor); foretibia possessing (metasomal 2) (Abd III) (Figure 3), wider than strigil (pectinate curvate spur) ventrodistally petiole, anterior foramen vertically set off, forming antenna cleaner with curvate and formed of first helcium (Bolton 1990),

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adjusted into posterior petiolar foramen, poststernite largest metasomal sclerites posttergite largest sclerite of postpetiole comprising approximately 3/4 of generally subfungiform (Figures 3, 9, 32, 34), opisthogastral region; posttergite convex posteriorly augmented forming postnodus (Figure 9) or slightly depressed [rudis]; (Figure 3), usually with anterolateral lobes spiracle anteriad, subdorsad. Metasomal 4 to [belti] posterolaterad to cinctus 2; postnodus pygidium somites non-specialized, pre- and usually composed of dorsal and posterodorsad posttergites differentiated by pronounced faces [scabriusculus, clathratus, rudis], carina (Longino and Snelling 2002); pygidium continuously convex posteriad (the two faces (Abd VII) divided into epipygium (tergite) not differentiated by postnodus) (Figure 31) and hypopygium (sternite) (Figures 3, 9). [convergens], or dorsally flat, narrowing into postnodus and somewhat vertically set off Gynes (Figures 14-16) forming lamella posteriorly [mayri, batesi]; Although similar to workers, gyne are larger tergite and sternite separated by dorsosternal with thoracic sclerites (Figures 15-16) Downloaded from sutures, fused (Bolton 2003); poststernite corresponding to alates in Apocritans. Ocelli crescentiform, emarginate posteriorly, leaving posteriad within frons (Figure 14). helcial metasomal 3 (Abd IV) presternite visible (Figures 9, 13, 16), projected Gyne variations on mesosoma (Figures 16, http://jinsectscience.oxfordjournals.org/ anteromedially into sternopostpetiolar 17). Mesonotum divided into anterior scutum (“subpostpetiolar”) process (Figure 9); and posterior scutellum by curvate, sternopostpetiolar process forming with scutoscutellar, or prescutellar groove; scutum cinctus 2 (Figure 13) transversal (trans- divided by transcutal suture into greater sternal) cavity ventrally, where petiolar anterior sclerite, and posterior prescutellar sternite posterior margin couples; region, dorsomesial between axillae; ventropostpetiolar process somewhat conic, parapsidal lines extended posteroanteriorly

transversally truncate (Figure 13), blunt from transcutal suture to discal scutum, by guest on September 29, 2015 [spiniperdus], unilobate [mayri] or bilobate slightly diverging anteriorly; axillae laterally apically; caudal postpetiolar foramen longer and wider, forming prescutellum and posteroventrad giving posteroventrad position embracing scutellum anteriorly; axillular (lateral view) to opisthogaster (metasomal 3 scrobe (“fossa”), where wings rest, lateral, [Abd IV] to pygidium, see Metasoma in under axilla, formed of lateromedially Materials and Methods) (Figure 3); impressed axillar groove, running from wing postpetiolar spiracle anteriad, slightly ventrad. axillar sclerites to scutellum; Metasomal 3 (Abd IV) (Figure 3), largest mesoanepisternum and metanepisternum metasomite, first opisthogastral somite, third separated by lateral mesometathoracic specialized (possessing second helcium and (mesometapleural) suture; metanepisternum third cinctus) metasomite occupying nearly and metakatepisternum separated by short 2/3 of metasoma; elliptical or ovate; metanapleural sulcus. Wings similar to those presclerites forming second helcium (Bolton of the male. 1990) (Figure 3); stridulatory organ (Wheeler 1984) formed between metasomal 2 and Sculpture (Figures 15, 17-34) second helcium; helcial sclerites and Sclerites usually exhibit combination of two postsclerites separated by cinctus 3 (“girdling or three sorts of sculpturing. Sculpture in constriction”) (Figures 3, 9); postergite and Procryptocerus divided into microsculpture

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(background sculpture), and macrosculpture attenuatus, convexus, carbonarius-posteriad], (regular [circular] or irregular depressions, or or finelly striate (Figures 15, 32) [some longitudinal and transverse elevations). Andean, Mesoamerican, and northern South Microsculpture: micropunctate, American species] [scabriusculus, tortuguero, microreticulate, microimbricate or marginatus, spiniperdus, ferreri]. Alveolate microstrigulate. Macrosculpture: impressed sculpture of Andean and Mesoamerican holes without costae, ridges or carinae species on frons posteriad, pronotum anteriad, (foveate, foveolate, punctate), at level of tergal petiole [eladio, FS_11] and postpetiole surface (shallow) (striolate, imbricate, (Figure 31); mesosoma, petiole and areolate, dotted, puncticulate), raised (costate, postpetiole porcate [batesi, mayri], tergal carinate, carinulate, vermiculate, striate, postpetiole rugocostate [Brazilian species] sulcate, strigate), or their combinations [regularis, sampaioi, convergens, schmalzi], (scrobiculate, porcate, alveolate, rimose). femora costulate or costate in Central Often, when integument smooth and polished American [paleatus, impressus] and Brazilian Downloaded from (shiny = glossy), dorsum, especially on [schmalzi] species. metasomal 3 sclerites, micropunctulate and bears combination of other micro sculpture. Clathrate sculpture on frons and promesonotum [carbonarius, rudis, batesi, http://jinsectscience.oxfordjournals.org/ Surfaces normally without sculpturing: mayri, clathratus] (Figures 21, 22); costate or torulus, hypostoma, funicular segments, and costulate sculpturing often on mandibles, postocciput. Surfaces regularly micropunctate, clypeus (nasus), nasal flanks, frons (Figures and without macrosculpture, neck, 14, 15, 18), malar space, temple, vertex prosternum, mesonotal lobes, ventral (Figure 15), gena, promesonotum, discal metepisternum, propodeal spines, lateropronotum, mesopleuron, propodeum, posteropropodeum, sternal petiole and coxae, femora, tibiae, and metasomal tergites

postpetiole, and metasomal 3 in Andean 1, 2, 3; rimae (ondulate striae or costae) more by guest on September 29, 2015 species over 600 m of elevation. Surfaces common on frons [sampaioi, victoris, microreticulate or microimbricate, without convergens] and mesonotum [victoris], when macrosculpture: scape (almost always metasomal tergite 3 punctulate (densely microimbricate), femora (microimbricate or punctate), some species have farinose texture microstriolate). Elevated ridges (costae and [impressus, belti, subpilosus]; scrobiculae carinae) often microsculptured (micropunctate often bordering areas as vertex [mayri, or microimbricate) on background. Striations schmitti, clathratus], temple [eladio], and sulcations more common in Brazilian lateropronotum posteriad [scabriusculus], species. Circular impressed sculptures and mesepisternum anteriad [scabriusculus], combination with costae, striations and notopropodeal fusion when grooved [schmitti, sulcations more common in Andean and coriarius], petiole and postpetiole posteriad, Mesoamerican species. cincti 2 and 3 (Figures 15, 32).

Frons clathrate in Andean species [mayri, Mandible often longitudinally costulate batesi] and Brazilian clathratus (Figures 21, (Figure 4); anteclypeal region often strigate, 22) or foveolate (schmitti, coriarius,nalini); discal clypeus ecarinate, variably metasomal 3 punctate [belti, impressus longitudinally costate, or with medial carina (Figure 33)], glossy [eladio, belti, mayri, or costa (Figures 3, 4), nasal flank ecarinate or

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costulate; clypeal carina often extending back refers to exceptionally short, fine hairs and continuing mesiad, parallel and very close forming second layer beneath pilosity to frontal carina; frontal carina describing (Hölldobler and Wilson 1990). more or less straight line [eladio], curvate [scabriusculus], convergens, regularis, Procryptocerus bear both short and long, subpilosus, coriarius] (Figures 3, 4, 14), flexous (flagellate) pilosity (Figure 26), or sinuate [belti] (Figure 20) or sinuate-bilobate short and long, stiff or subspatulate pilosity [mayri, batesi]; frons foveolate (or foveate) (Figures 22, 30, 31); longest pilosity on tergal [nalini, eladio, pictipes-anteriad] (Figure 23), petiole and postpetiole [mayri, impressus,]; foveate-costulate [scabriusculus] (Figure 14), often medial dorsal line of meso and striate [adlerzi] (Figure 17), costulate metasoma denudate; lateral hairs of [virgatus], costate [regularis], reticulate or anteclypeus pecten (Figure 22) commonly areolate [belti, hirsutus, convexus, pictipes] convergent; frons more common with stiff, (Figure 20), reticulate [pictipes-anteriad], scattered pilosity; frons posteriad and Downloaded from areolate [pictipes-posteriad], clathrate and frontovertexal margin usually with two areolate [batesi] (Figure 21), infra transverse lines of uniformly separated stiff lateropronotum porcate or costate [most hairs slightly directed anteriorly; eyes species] (Figure 29); mesopleuron porcate or denudate; malar space often possessing few http://jinsectscience.oxfordjournals.org/ costulate (Figure 29), and foveolate; scattered hairs; shorter flexous pilosity or propodeal spines microsculptured or ecarinate pubescence on postgenal bridge; mandible and glossy, posteropropodeum supra strigate with erect short hairs, ventrally and distally [hylaeus, mayri], ecarinate and shiny [mayri], with flexous pilosity; scape usually with short, or striate (or longitudinally costate) stiff, uniformly distributed, sparse hairs [montanus, striatus]; meso and metapleura (Figure 22) promesonotum and usually costulate (Figure 29); nodal truncation dorsopropodeum with long stiff or flexous

ecarinate and glossy [belti] or strigate hairs; propleuron denudate or pubescent; meso by guest on September 29, 2015 [scabriusculus]; tergal petiole and postpetiole and metapleura usually denudate; coxae both areolate [coriarius] (Figure 31); sternal petiole pubescent and with flexous hairs on ventral, and postpetiole ecarinate [most species]; dorsal and posterior faces, usually medially metasomal 3 tergite striate-costate (Figure pubescent combined with some flexous, 15), costulate, or costate [several species in sparse hairs; dorsum of femora and tibiae the whole range of the geographical usually with decumbent and subdecumbent, distribution] (Figures 15, 16, 32); metasomals stiff hairs; petiole and postpetiole 4-6 pretergites smooth, postergites strigulate dorsolaterally with subdecumbent or suberect or microtuberculate; epipygium punctulate; stiff or flexous hairs, ventrally usually metasomal 3 sternite ecarinate [most species], denudate; ventropostpetiolar process denudate striate (or costulate) [ferreri] (Figure 34). or bearing few scattered, long, flexous hairs, rarely pubescent, metasomal 3 (Abd IV) Vestiture (Figures 22, 26, 30, 31, 35) tergite denudate or bearing either erect or Two kinds of vestiture are present in suberect, long or short, flexous pilosity, or Procryptocerus. Pilosity (Figures 26, 30, 31) combination of both; exception of pubescent refers to long, erect, suberect, subdecumbent, metasomal 3 tergite is Andean species from decumbent (Figure 35), or appressed (not Panama and Venezuela; ventral metasomal 3 drawn) hairs, and pubescence (not drawn) denudate, pubescent, or with combination of

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pubescent and short, scattered, flexous hairs; Head (Figures 36-38) metasomals 4 to pygidial postergites with few Subglobular, never transverse. Interocular erect or suberect flexous or stiff hairs, often distance shorter than, or subequal to, median arranged in transverse lines with shorter hairs head length; mandibles curvate mesially; [species at low elevations]; hypopygium anteclypeal carina medially weakly notched; pubescent or not, and often possessing short, clypeus protruded into nasus; frontoclypeal subdecumbent, flexous hairs. Brazilian sutures modified in transverse groove between species usually with stiff pilosity shorter than toruli; frontal carina short, divergent caudad; flexous hairs of some Andean and vestigial to obsolete behind eyes; antennal Mesoamerican species, short stiff hairs in scrobe above eyes; vertex not distinctly Andean convexus, hirsutus, belti. Some deflexed; frons posterior corners distinct to Andean species may bear flexous or stiff, long obsolete. Eyes lateral, strikingly protruding, or short pilosity; other species may possess slightly extending dorsad and ventrad, abundant flexous pilosity [batesi]. Some comprising most of head; ocelli protruding in Downloaded from species almost denudate dorsally and most species dorsally, posteriad to eyes, pubescent ventrally, especially on anteriad to vertexal margin, sometimes opisthogastral sternites [eladio]. Few assemblaged on ocellar triangle; antenna Brazilian species with some very scarce long filiform, 13-segmented; scape subequal to or http://jinsectscience.oxfordjournals.org/ flexous hairs on caudal metasoma posttergites. longer than second funicular segment.

Color Mesosoma (Figures 37, 38) Color usually varies from dark-orange or red- Trunk-shaped, humped; scutum with deeply brown appendages, and black meson in some impressed notauli; anterior branches longer Andean species usually over 600 m of than the posterior medial stem; episternum elevation, to completely black in most species superior to mesocoxa, usually ecarinate;

found at low elevations. Minor color dorsopropodeum with small, spiniform by guest on September 29, 2015 variations are as follows. Scape and pedicel process posterolaterally; femora moderately yellow, orange, red, or brown; eye brown; concave mesially and incrassate in middle; all palps yellow; mandible brown laterodistad; segments of legs comparatively long and flagellum, tibiae and telotarsi orange-brown; slender; middle and hind tibiae usually remaining body black. without apical spur.

Male (Figures 36-45) Metasoma (Figures 37, 38) Male longer, slender than gyne (Figure 37), Metasomal 1 (Abd II = petiole) sessile, ranging from 4.8 mm [COL] to 9.9 mm elongate, subcylindrical. Metasomal 2 (Abd [scabriusculus]. Following traits separate III = postptiole) similar to metasomal 1, Procryptocerus males from others in the Tribe somewhat shorter, more incrassate posteriad; Cephalotini: scape long, subequal to or longer metasomal 3 (Abd IV) largest metasomal than second funicular segment; postpetiole somite, longer than petiole and post-petiole longer than height; mandibles strongly combined; hypopygium rounded posteriad mesially curvate; head subglobular; (Figure 40) [scabriusculus], truncate [batesi], posterolateral spines or teeth on subtriangular or conic [adlerzi]; paramere dorsopropodeum (Kempf 1951). (Figures 41-43) paddle-shaped, rounded apically, sometimes about same length of

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caudal metasoma ; volsella (Figure 45) mesiad to paramere, bifurcate into cuspis and digitus Along with different shapes of discoidal and volsellaris; cuspis volsellaris mesial to first submarginal cells on anterior wings, main paramere, subcylindrical, sinuate, shorter than characters that separate species are variations digitus, truncate apically; digitus volsellaris within external and internal genitalia, which mesial to cuspis, compressed, hook-shaped, contain well-developed hypopygium, ventrally bent distad. volsellae, and parameres (Kempf 1951).

Wings (Figure 46) Discussion Wing shape, venation and cells similar in both male and gyne; male fore wing extending to Torulus vs. Annulus (antennalis) (Figures 4, level of posteriad caudal metasoma; hyaline 14) [goeldii], or infumate [batesi, mayri, Gauld and Bolton (1988) consider the torulus impressus, scabriusculus]. Useful, distinct to be the socket, or the cephalic foramen, in Downloaded from variations have not been found for which the antennal condylar bulb inserts. discriminating species within Procryptocerus. Bolton (1994) considers torulus to be the Fore wings with distinct anterodistad stigma; small annular sclerite that surrounds the in anteroposterior sequence, proximal antennal socket. Torre-Bueno (1989) http://jinsectscience.oxfordjournals.org/ (proximad to stigma) longitudinal veins are C considers the annulus (antennalis) to be the (Costa), Sc+R+Rs ring sclerite of the head into which the basal (Subcosta+Radial+Radiosectorial), M+Cu segment of the antenna is inserted. Gordh and (Medial+Cubital), and A (Anal). Distad Headrick (2002) consider annulus to be the longitudinal veins are R, M and Cu; distad antennal sclerite forming a sclerotized ring on veins do not reach apex of wing; recurrent the head into which the basal segment (scape) veins are cu-a (cubital-anal) and m-cu of the antenna is inserted. We follow Bolton

(medial-cubital); cells formed by confluence (1994). by guest on September 29, 2015 between longitudinal veins or between longitudinal and recurrent veins; proximally, Mesosoma (3, 9) three cells present: Costal (CC), Basal (BC) or For the second tagma, the term “alitrunk” (ali Radial (“Media”), and Subbasal (SBC) or = wings) has been proposed to avoid Cubital (“Submedia”); posteriad to stigma, confusion with “thorax” (Gauld and Bolton Submarginal-one cell (SMC1) (closed) and 1988, Bolton 1994, Wilson 2003). Submarginal-two cell (SMC2) (opened) are Nevertheless, a similar confusion could occur present; Discal cell-one (DC1) posteriad to between metasoma and “abdomen”. Submarginal-one and distad to Basal cell (or “Alitrunk” is not recommended since worker Radial cell), formed by confluence of M, Cu, ants do not possess wings, and mesosoma is Rs and m-cu. Distal field without cells. Hind well characterized in Apocrita (see Nauman wing possessing proximally two distinct 1991). longitudinal veins: R+Rs and M+Cu; cu-a is basad in proximal field; Basal cell BC (or Notopropodeal fusion Radial cell) closed distally by M vein; The mesosoma comprises the thorax plus the Subbasal SBC cell (or Cubital cell) closed propodeum, the tergite of Abd I fused to the distally by cu-a and posteriorly by 1A; distal thorax. The pleurites and the sternite of Abd I field without distinct veins. are entirely reduced and the tergite remains. In

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workers of ants, the notal and propodeal covers the background of the cuticle. sclerites are usually fused forming a tergal Microsculpture may be present on the (notal) fusion between the notum and elevations or impression of the propodeum. This condition is a notopropodeal macrosculpture, usually when the fusion. Externally, it is usually impossible to macrosculpture (e.g. costae, striae) is glossy recognize the structures involved in the (shiny), or, more commonly, in smooth fusion. The line of fusion may be indistinct surfaces devoid of macrosculpturing. (notopropodeal fusion usually convex), Micropunctulae and microimbricae are fairly obsolete or differently marked by a suture, different, but require close inspection to groove, impression, depression, etc. The line interpret. The micropunctulae condition of fusion has different names in the literature, occurs as microscopic pricks more common such as “propodeal suture” (a suture on the on opaque surfaces, whereas microimbricae propodeum), “metanotal suture” (a suture on are either overlapping microscales (as tiles on the metanotum), “metanotal groove” (a a roof) or microreticulae (“dotted”) that give Downloaded from groove on the metanotum), “metanotal an overlapped appearance. The latter are more impression” (an impression on the common on glossy surfaces. Macrosculpture metanotum), “metapropodeal suture” (a suture can be divided into impressed, superficial, on the posterior [meta] region of propodeum), raised and combined sculpturing with the http://jinsectscience.oxfordjournals.org/ antepropodeal suture, metanotal area, etc. following states. 1. Impressed macro Since these terms make reference to the line sculpture: circular or oblong (punctate, of fusion, we recommend using the adjective foveate, foveolate); linear (furrow-like) notopropodeal in reference to the line of (furrowed, grooved, sulcate). 2. Superficial fusion; for instance: notopropodeal suture, macro sculpture (non-impressed cuticle, notopropodeal groove (Figure 3), surface spaces between costae or carinae): notopropodeal convexity, notopropodeal linear spaces delimited by costae or ridges -

impression, notopropodeal excavation, or costae or ridges are low and same width as by guest on September 29, 2015 otherwise make reference to the striae- (striate, striolate, strigate, strigulate); notopropodeal fusion to describe specific non-linear spaces delimited by costae or characters, such as notopropodeal fusion flat, ridges (reticulate, areolate). 3. Raised macro notopropodeal fusion convex, etc. In several sculpture (carinate, carinulate, costate). 4. groups, e.g. some Camponotus, the Combinations between impressed and raised metanotum is clear and so are the macrosculpture: canalicular (porcate, mesometanotal suture and the metanotal- scrobiculate); polygonal and irregularly propodeal suture. In those cases, a polygonal (alveolate), or irregular porcae, notopropodeal fusion is not apparent. irregular reticulae, irregular rugocostae- Occasionally, a mesometanotal fusion or a alveolate (clathrate). metanotal-propodeal fusion can also be identified in a very few cases. Striate sculpturing refers to longitudinal lines on a non-impressed cuticle, running parallel Sculpture (Figures 14-34) between thin and low elevated costae or Procryptocerus species bear both micro- and costulae; the costae (or costulae) and lines are macrosculpture. Microsculpture narrow and about the same width. The (micropunctulate, micropunctate, sculpture should be named striate-costulate, microimbricate, microstrigate, “dotted”) but it is customary to call it striate. Striate is

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one of the most common forms of sculpturing porcate when a set of combinations of costae in Procryptocerus; it is often present on the and impressions between costae are present, metasomal 3 (first opisthogastral) tergite. forming canalicular (sulcal) spaces. Costate sculpturing refers to costae (elevated Anastomosed porcae are porcae that run into ridges rounded at their edges, dim. costulae) each other. Scrobiculate are surfaces where in general running parallel or quasi parallel to scrobiculae (parallel, short porcae) are each other, the interspaces are wider than uniformly organized in a contiguous, chain- costae. Rimose refers to nearly parallel like series. When there are septae within excavations (rimae), often narrow, short or striae, the sculpturing is reticulate (quadri- or long, in the shape of wavy cracks, running quasi quadriculate) [belti frons] or areolate into each other (Gordh and Headrick 2000); (polygonal or quasi polygonal) [scabriusculus elevations between rimae are vermiculate, frons posteriad]. In a subsecuent stage there often wide and flat at their ridges, which are are septae within a porcate surface, and the usually micropunctulate. Rimae refer to the sculpturing is alveolate or clathrate. Alveolae Downloaded from longitudinal fissures, crack-shaped are lacunose, impressed spaces delimited by interspaces; ridges refer to the elongated irregular rugae or “costae” with sharp rims at elevations (costae). The costae are wavy. The their edges. The alveolae are regular or combination between rimae and flat ridges, irregular polygons, and the sculpturing is http://jinsectscience.oxfordjournals.org/ running in anastomosis, produces rimose called alveolate. Alveolate cuticle is often (dim. rimulose) or rivose (dim. rivulose) present on the posterior frons and petiole. sculpturing. Combinations of character states, Surfaces having alveolate sculpturing in shuch as rimose-vermiculate, or rivose- Procryptocerus form landscapes of lacunae vermilaculate could be more specific. between ridges (ranges of “hills”) containing sharp or obtuse edges. Clathrate sculpturing Punctures are slightly impressed points refers to the most complex, irregular

(pricks) on the cuticle that appear to be made combination of irregular porcae and by guest on September 29, 2015 by a needle (Gordh and Headrick 2000). transversal septae (crossing costae), forming Punctures constitute the smallest circular- deep, alveolate, cancelled holes of different macrosculpture. Derived adjectives describing and irregular diameter. In clathrate sculpture, this sculpture are punctate (with punctures), the costae run anteroposteriorly in irregular puncticulate (sparcely punctate), punctulate fashion prevailing over the short transverse (closely punctate). When puncticulate and costae forming the septae. High density of punctulate are present on the same surface, the alveolae conform clathrate sculpture. This difference between the sculptures is clear. characteristic is the most important one to When only one is present, the terms are differentiate clathrate and reticulate interchangeable. When densely punctate, the sculpturing, which could be apparently similar cuticle has a farinose texture. Dots (dotted when both are present on frons and when sculpturing) are non-impressed circular marks, looking at them through a common they are superficial, rounded, and the same microscope. Clathrate sculpture may be size as micropunctures. A costate integument present on the frons and mesonotum [mayri, emphasizes the costae and not the interspaces batesi, clathratus]. Reticulate sculpture is (striae or sulci); in these cases spaces between usually present on frons only [belti, convexus, costae are in general wider than the costa hirsutus]. edges and not impressed. The sculpture is

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When a combination of sculptures is present, it is useful to hyphenate two, sometimes three, Sculpturing within Procryptocerus is a rich different words qualifying sculpture (e.g. source of characters, which is helpful in rugo-costate, costate-foveate, foveate- stablizing and recognizing the identity of reticulate, micro-striolate-imbricate). In species and could permit the formalization of general, when differences between hypotheses of evolutionary trends. proportions occur, the first word should emphasize the most common sculpture, or emphasize the first sculpture when referring to an anteroposterior (or any directionality) sequence of the sculpture present on any surface. Discriminating thickness of raised sculpture (e.g. costate, carinate) and width of circular, impressed sculpture (e.g. punctate, Downloaded from foveate, foveolate) is often not clear when only one of these types of sculpture is present. In those cases, the closest terms might be used interchangeably (e.g. foveate or foveolate, http://jinsectscience.oxfordjournals.org/ costate or costulate, rimose or rimulose). Discrimination of those sculptures is easier when several types of sculpture are present in the same area of an ant.

Confusion occurs between sculpture texture (appearance) and sculpture structure

(constitution). To recognize the constitution by guest on September 29, 2015 (nature) of the sculpture, textures (e.g. leathery, farinose, rugous, coriarious, corticinus, etc.) should be avoided. Appearance strongly depends upon the “momentary” criterion of the researcher or interpreter, and magnification, light or system (microscope, SEM images, photomontage) used to recognize it. Nonetheless, when using a common microscope or photomontage images, the appearance is sometimes quite distinct with some descriptive forms (e.g. politus, shiny, glossy, farinose). It is best to use SEM images of sculpture (For instance see http://www.evergreen.edu/ants/genera/Antsof CostaRica.html). On the other hand, drawings are the best way to convey information about boundaries of sclerites.

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Acknowledgments Donalt Agosti (AMNH), Gary Alpert and Stefan P. Cover (MCZ), Patricia Rojas (Most codens standing for collections can be (Instituto Ecologia Xalapa - Mex.), Israel del found in http://hbs.bishopmuseum.org/ Toro, Melissa Ferkie and The Department of codens/codens-inst.html) Entomology of the University of Arkansas, To Erika Valentina Vergara (UNAB) for her Clara Nichols (The University of California at permanent support during this project and Berkely), Pacheco-Lucero family, Ayda making important suggestions to the Navarro, Eduardo Amat, Raúl Vélez, Adolfo manuscript. Barry Bolton, Phil Ward, Cesare Molina, Alejandro Madrigal, and Sandra Baroni-Urbani, and Jack Longino reviewed Uribe. Other financial support comes from the manuscript and made important Unal-Colciencias-Laspau agreement, The suggestions specially to the treatment of the University of Texas at El Paso (UTEP), The metasoma. Carl S. Lieb reviewed the Dodson Fellowship UTEP, Idea Wild, manuscript. Other researchers and institutions Universidad Nacional de Colombia (UN), and Downloaded from have supported and loaned specimens to this Juan G. Serna-Cardona. Project, including Celsa Garcia (UNAB), Phil Ward (CIPW), Jack Longino (Evergreen State Editor’s note: Paper copies of this article College), Jeffrey Sossa, Ted Schultz, and will be deposited in the following libraries. http://jinsectscience.oxfordjournals.org/ David Furth (USNM), E. Richard Hoebeke The date of publication is given in ‘About the and Dimitri Forero (CUIC), Roberto Poggi, Journal’ on the JIS website. and Fabio Penati (MSNG), Doug Yanega Universitaetsbibliothek Johann Christian (UCRC), Weiping Xie (LACM), Bernhard Senckenberg, Frankfurt Germany; National Merz (MNHG), Doug Yanega (UCRC), Museum of Natural History, Paris, France; Michael Thomas (FSCA), Mónica Ospina, Field Museum of Natural History, Chicago, and Diego Perico (IAvH), David Donoso Illinois USA; University of Wisconsin,

(QCAZ), Kim Goodger, and Conrad Gillett Madison, USA; University of Arizona, by guest on September 29, 2015 (BMNH), Carlos Brandao, Rodrigo Feitosa, Tucson, Arizona USA; Smithsonian Rogerio Rosa-Dasilva, Flávia A. Esteves, and Institution Libraries, Washington D.C. USA; Cristiane P. Scott-Santos (MZSP), John Lattke The Linnean Society, London, England. (MIZA), Takumasa Kondo (UCD), Enrique Torres, David Cuéllar, and Teresa Mosquera References (UN), José Palacios-Vargas (UNAM), Javier Martínez (UNAB), Martha Penaranda, Agosti D, Majer JD, Alonso LE, Schultz T. Norelhy Quimbayo (UNAB), Gabriela 2000. Ants. Standard Methods for Measuring Castano (UNAM), John Quiroz (UNCM), and Monitoring . Smithsonian Stefan Schödl (NMW), Daniel Burckhardt Institution Press. Washington. (NHMB), Alex Wild and his webpage http://www.myrmecos.net, Manuel Solís Bohart RM, Menke AS. 1976. Sphecids (INBio), Antonio Mayhé-Nunez (CECL), Ana Wasps of the World. A Generic Revision. Yoshi Harada (MPEG), Orlando Tobias University of California Press. Berkeley. Silveira (MPEG), Giselle Zambrano (Universidad del Cauca - Col.), Patricia Bohart RM, Stange LA. 1965. A Revision of Chacón (Universidad del Valle – Col.), Zulma the Genus Zethus Fabricius in the Western Gil (CENICAFÉ - Col.), Bryan Fisher (CAS), Hemisphere (Hymenoptera: Eumenidae).

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University of California Publications in Emery C. 1887. Catalogo delle formiche Entomology, Vol. 40. esistenti nelle collezioni del Museo Civico di Genova. Parte terza. Formiche della regione Bolton B. 1990. Abdominal characters and Indo-Malese e delláustralia. Annali del Museo status of the Cerapachyine ants (Hymenopera, Civico di Storia Naturale di Giacomo Doria Formicidae). Journal of Natural History 24: 4: 209–258, 3 Figures, 2 plates; 5: 427–473, 2 53-68. plates.

Bolton B. 1994. Identification Guide to the Emery 1924 (1922). Hymenoptera. Fam. Ant Genera of the World. Harvard University Formicidae. Subfam. Myrmicinae. In: P. Press. Wytsman, editor, Genera Insectorum, fasc. 174C: 207-397. Bolton B. 2003. Synopsis and Classification of Formicidae. Memoirs of the American Gauld ID, Bolton B. 1988. The Hymenoptera. Downloaded from Entomological Institute. 71: 1-369. The University Printing House. University of Oxford.

Bolton B. 2007. How to conduct large-scale http://jinsectscience.oxfordjournals.org/ taxonomic revisions in Formicidae, pp. 52-71. Gordh G, Headrick DH. 2001. A Dictionary of In: RR Snelling, BL Fisher, PS Ward, editors. Entomology. CAB International Publishing. Advances In Ant Systematics (Hymenoptera: Formicidae): Homage To E. O. Wilson – 50 Harris KA. 1979. A Glossary of Surface Years Of Contributions. Memoirs of the Sculpturing. University of California, American Entomological Institute, 80. Department of Food and Agriculture, Division of Plant Industry, Laboratory Services. Bolton B, Alpert G, Ward PS, Nastkrecki P. Occasional Papers in Entomology, No. 28. 2006. Bolton’s Catalogue of Ants of the by guest on September 29, 2015 World: 1758-2005. Harvard University Press. Hölldobler B, Wilson EO. 1990. The ants. (CD available). Harvard University Press.

Brown RW. 1979. Composition of Scientific Kempf WW. 1951. A taxonomic study on the Words. Smthsonian Institution Press. ant tribe Cephalotini (Hymenoptera: Formicidae). Revista de Entomologia 22:1- Baroni Urbani C, de Andrade ML. 2003. The 244. ant genus Proceratium in the extant and fossil record (Hymenoptera: Formicidae). Museo Kempf WW. 1957. Sôbre algumas espécies de Regionale di Sciénze Naturali Monografie 36: Procryptocerus com a descrição duma espécie 1-492. nova (Hymenoptera, Formicidae). Revista Brasileira de Biologia 17: 395-404. de Andrade ML, Baroni Urbani C. 1999. Diversity and Adaptation in the Ant Genus Kukalová-Peck, J. 1991. Fossil History and Cephalotes, Past and Present. Stuttgarter the Evolution of Hexapod Structures. (Chapter Beitrage zur Naturkunde. Serie B (Geologie 6), pp. 141-179. In: Naumann ID, Carne PB, und Palaontologie), Nr. 271. Lawrence JF, Nielsen ES, Spradbery JP, Taylor RW, Whitten MJ, Littlejohn MJ,

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editors. The of Australia. Volume 2. ouvrières. Contribution à la systématique et à CSIRO. la phylogénie des fourmis (Hymenoptera : Formicidae). Annales Société entomologique Longino, J.T. 2006. de France 40 (3-4): 291-371. http://academic.evergreen.edu/projects/ants/A ntsofCostaRica.html. The Evergreen State Schultz TR. Alonso LE. 2000. Glossary. Pp. College, Olympia WA. 223-229. In: D Agosti, JD Majer, LL Alonso, TR Schultz, Editors. Ants. Standard Methods Longino JT, Snelling RR. 2002. A for Measuring and Monitoring Biodiversity. Taxonomic revision of the Procryptocerus Smithsonian Institution Press. (Hymenoptera: Formicidae) of Central America. Contributions in Science, Number Snodgrass RE. 1993. Principles of Insect 495. Natural History Museum, Los Angeles. Morphology. Cornell University Press. Downloaded from Mackay WP, Vinson SB. 1989. A Guide to Sparks SD. 1941. Surface anatomy of ants. species identification of New World Ants Annals of the Entomological Society of (Hymenoptera: Formicidae). Sociobiology, 16 America 34: 572-577. (1):1-47. http://jinsectscience.oxfordjournals.org/ Triplehorn CA, Johnson NF. 2005. Borror and Mackay WP, Mackay EE. 2003. The ants of Delong’s Introduction to the study of insects New Mexico (Hymenoptera: Formicidae). The 7th edition. Brooks/Cole Beltmont. Edwin Mellen Press. Ward PS. 1999. Systematics, biogeography Mackay WP, Mackay EE. 2006. A new and host plant associations of the species of the ant genus Pachycondyla F. Pseudomyrmex viduus group (Hymenoptera:

Smith, 1858 from Ecuador (Hymenoptera: Formicidae), Triplaris- and Tachigalia- by guest on September 29, 2015 Formicidae). Myrmecologische Nachrichten inhabiting ants. Zoological Journal of the 8:49-51. Linnean Society 126:451-540.

Naumann ID. 1991. Hymenoptera (Wasps, Ward PS. 2004. Ant Morphology. The Ant bees, ants, sawflies). pp. 916-1000. In: ID Course No.4. August 2004, Costa Rica. Naumann, PB Carne, JF Lawrence, ES University of California at Davis. Entomology Nielsen, JP Spradbery, RW Taylor, MJ Department. 4 pp. (unpublished). Whitten, MJ Littlejohn, editors, The Insects of Australia. Second Edition. Volume 2. CSIRO. Ward PS, Downie DA. 2005. The ant subfamily Pseudomyrmecinae (Hymenoptera: Torre-Bueno JR de la. 1989. The Torre-Bueno Formicidae): phylogeny and evolution of big- Glossary of Entomology, compiled by SW eyed arboreal ants. Systematic Entomology 30, Nichols, NewYork Entomological Society. 310–335. American Museum of Natural History. New York. Ward PS 2007. Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Perrault GH. 2004. Étude morphoanatomique Formicidae). Pages 549-563 In: ZQ Zhang, et biométrique du métasoma antérieur des WA Shear. Linnaeus tercentenary: Progress in

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invertebrate taxonomy. Zootaxa 1668. Magnolia Press, Auckland. 766 pp.

Wheeler DE. 1984. Behavior of the ant, Procryptocerus scabriusculus (Hymenoptera: Formicidae), with comparisons to other cephalotines. Psyche 91:171– 192.

Wheeler QD. 2008. Undisciplined thinking: morphology and Henning’s unfinished revolution. Systematic Entomology 33: 2-7.

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Figure 1. Mushroom-shaped proventricular valves found in the tribe Cephalotini (Redrawn from Emery 1922). High quality figures are available online.

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Figure 2. Dorsal (upper), and lateral (lower) representation of a hypothetical insect showing common positions and orientations used to discribe bilateral organisms. High quality figures are available online.

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Figure 3. Procryptocerus scabriusculus. Dorsal view. High quality figures are available online.

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Figure 4. Procryptocerus scabriusculus. Worker. Head, dorsoanterior (frontal) view. Above: face. Lower: half side of face zoomed (part) to show morphological details. High quality figures are available online.

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Figure 5-8. Procryptocerus scabriusculus. Worker. Buccal pieces. 5-7 right mandible: 5: dorsal, 6: lateral, and 7: mesial views; 8: posterior view of maxilolabial complex. High quality figures are available online.

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Figure 9-10. Procryptocerus scabriusculus Worker. Lateral (profile) view; 10: Procryptocerus scabriusculus. Worker. Mesosoma profile zoomed to show details. High quality figures are available online.

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Figure 11-13. Worker. Mesosoma and waist region. Ventral view. High quality figures are available online.

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Figure 14. Gyne. P. scabriusculus. Head frontal view. High quality figures are available online.

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Figure 15. Gyne. P. scabriusculus. Dorsal view. High quality figures are available online.

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Figure 16. Gyne. P. scabriusculus. Lateral (profile) view (habitus). High quality figures are available online.

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Figure 17-25. Sculpture on head. 17-23 frontal (dorsoanterior); 24 lateral (profile); 25 dorsoposterior. 17 costate, 18 costate, concentricus in middle, 19 diverging costate posteriad, concentricus anteriad, 20 reticulate, 21 clathrate posteriad and in middle, rugocostate anterolaterad, anastomosate anteromesiad, 22 clathrate, 23 foveate or foveolate, 24 parietal costate or rugocostate, frons clathrate, 25 vertex strigate. High quality figures are available online.

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Figure 26-34. Sculpture on pro, meso and metasoma. 26 vertex striate (or longitudinally costate), mesosoma costate- vermiculate (or rimosus), 27 vertex strigate (or transversally costate), mesosoma costate, 28 (based on Longino 2006): promesonotum foveate, dorsopropodeum costate-porcate, 29 costate-sulcate, 30 (based on Longino 2006): lateropropropodeum supra foveate, infra costate, pleuron rugocostate, 31 (based on Longino 2006): rugocostate-alveolate, 32 petiole rugocostate, cinctus 2 scrobiculate, postpetiole costate (or porcate when interspaces are deep), metasomal 3 (Abd IV) striate-puncticulate (densely punctate), 33 metasomal 3 (Abd IV) tergite punctate, 34 metasomal 3 (Abd IV) sternite costulate- concentricus, glossy in middle. High quality figures are available online.

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Figure 35-38. General orientation of hairs. 36: Procryptocerus scabriusculus. Male. Head. Frontal (anterodorsal) view. 37: Procryptocerus scabriusculus. Male. Dorsal view. 38: Procryptocerus scabriusculus. Male. Lateral view; mtm: metasomal. High quality figures are available online.

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Figure 39-45. Procryptocerus scabriusculus. Terminalia. 39: epipygium; 40: hypopygium; 41, 42, 43: genitalia, 41: lateral, 42: dorsal, 43: ventral; 44: aedeagus; 45: volsella. High quality figures are available online.

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Figure 46-47. Male wings. Veins: C (Costa), Sc+R+Rs (Subcosta+Radial+Radiosectorial), M+Cu (Medial+Cubital), A (Anal), cu-a (cubital-anal), m-cu (medial-cubital). Cells: CC (Costal Cell), BC (Basal Cell) (or Radial), SBC (Subbasal Cell) (or Cubital), SMC1 (Submarginal Cell 1), SMC2 (Submarginal Cell 2), DC1 (Discal Cell 1), DC2 (Discal Cell 2); st: stigma. 47: Male legs. Above: foreleg; lower left: middle leg; lower right: hind leg (vestiture not drawn). High quality figures are available online.

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