Seven new species of Homeomastax are described, two from Panama and five from Costa Rica; a further new species from Colombia is noted, but not described owing to the lack of a male. The genus now contains 22 species. On the basis of the male genitalia the genus can be divided into 4 species groups, represented by the species surda, bouvieri, kressi and silvicola respectively; those species known only from the female (dentata, richteri, annulipes, and the new Colombian form) cannot yet be arranged in this scheme. Apart from male phallic characters, the species are externally recognizable only by the male and female subgenital plates and by morphometric ratios, especially E/F (elytron: femur). Of the three species redescribed by Rehn & Rehn (1934), the originally missing male of surda Burr and the female of veraguae Hebard are shown to have been correctly assigned by those authors. Their redescription of H. dentata Saussure, however, is invalid, as the type is a different taxon. The phallic complex of the genus is described in detail, and observations on copulation and the nature of the spermatophore are presented. We suggest that the hooked and tapered end of the spermatophore may be used as an intromittent organ by the copulating male.
The natural history and geographical distribution of the genus is reviewed; it occurs west and north of the Andean chains, though it extends up the major inter-Andean valleys of Northern Colombia, and ranges from Ecuador to Honduras. The four species groups are largely disjunct geographically. The bouvieri-group is South American, with an outlying species (brachyptera) in Central Panama; the surda-group is confined to the Pacific slope, the kressi-group to the Caribbean slope, and the silvicola-group is known only from the Caribbean slope and mountains of northern Costa Rica and southern Nicaragua.
Introduction
The Eumastacinae are a Neotropical subfamily of the American family Eumastacidae which, together with the Australian Morabidae and Biroellidae, constitute the Stenophalli, one of the four main divisions of the Eumastacoidea erected by Descamps (1973a). The validity of the clade Stenophalli is independently confirmed by molecular systematic techniques (Matt 1998). The great majority of the Eumastacinae, which currently include 11 genera, are South American; to date, only the genus Homeomastax has been recorded in Central America.
Homeomastax was split from Eumastax Burr 1899 by Descamps in 1979, primarily on the basis of the absence of an external pallium and differences in color, especially the red-tipped hind femora of Homeomastax. Previously (1971) the same author had recognized the taxon as the “surda species-group” of Eumastax, a usage different from that of Rehn & Rehn (1934), who indicated with this term a smaller subset of species within what is now Homeomastax.
The genus Homeomastax currently includes 14 species, which extend from the Pacific slopes of Ecuador and Colombia to Panama, Costa Rica, Nicaragua and Honduras. In Colombia the genus also extends from the north coast up the river valleys (Magdalena, Cauca) which run south between the different Cordillera of the Andes. On the other side of these barriers it is replaced by the closely related genus Eumastax. It is a wet-forest genus, and its members are most often found on ferns (e.g. Nephelea, Pteris, Hypolepis) which are, or at least can be, eaten by all species (see e.g., Descamps 1971: 109 and Rowell et al. 1983); however, they also eat many other plants, including some tree-ferns (Cyathaceae), Costus (Zingiberaceae), and Schistocarpha (Asteraceae) (pers. obs.), Piper (Piperaceae) and members of the Rubiaceae and Flacourtiaceae (Marquis 1991), and seedlings of trees, including Pentaclethra (Mimosaceae) (H.E. Braker, pers. comm.). The larvae are cryptically colored, dull green with olive-brown bands and speckles; the adults are brightly colored in shades of metallic blue, green, red and yellow, with black and white eyes. The general appearance is shown in Fig. 1. H. surda has been obtained from the canopy by insecticidal fogging (label data from specimens collected by H.R. Roberts in the collection of the ANSP). Unlike most forest grasshoppers, however, they are not confined to light gaps and/or the canopy, and larval groups especially can be found on shaded ferns in deep forest. Homeomastax species, like most eumastacids, are instantly recognizable by their habitual posture, with spread and depressed hind femora. The eggs are relatively enormous, more than half as long as the abdomen of the female (typically 9 X 2 mm each), and are matured only 2–6 at a time. The saw-like ovipositor valves suggest endophytic oviposition (see Braker 1989), but there is no direct evidence for this.
All the known species are colored very similarly, and differ externally only in size, the relative length of the wings, and subtle differences in the subgenital plates of both sexes. Their ecology also seems to be very similar. As far as is known, all species are allopatric. They are virtually impossible to identify to species in the field other than by distribution; usually a microscope is required, and sometimes a genital dissection.
The genus Eumastax (which then included Homeomastax) was reviewed by Descamps in 1971. In the present article we reexamine the latter genus. The initial motivation was the discovery of undescribed species in Central America; from our recently collected material we here describe seven new species, five from Costa Rica and two from Panama. We also report the existence of a further new species in the Colombian Chocó, so far represented only by females and therefore not formally described here. Previous descriptions have been based principally on the external genitalia, especially the shapes of the subgenital plates of both sexes. We supplement these with descriptions of the internal genitalia: those of the males (but not the females) present good taxonomic characters at the level of the species. In the course of this work we realized that the genus falls into four well-defined groups on the basis of the male genitalia, which has caused us to reexamine all the species for which the male is known. (H. dentata Saussure, H. annulipes Descamps and H. richteri Descamps are known only from the female, and we cannot determine therefore to which group they belong).
In a companion paper (Rowell & Bentos-Pereira 2001 p.255–262, this issue) we further describe from Panama two species of a new genus of eumastacine, Helicomastax, which is externally almost identical to Homeomastax, but which differs profoundly in the structure of the male internal genitalia.
Materials and methods
We used both pinned and alcohol specimens, the latter collected in the field into 70% alcohol to minimize distortion of the terminalia due to drying. In practice we found no difference in morphology between these and rehydrated pinned specimens. Phallic and spermathecal preparations were dissected, cleared in potassium hydroxide solution and lightly stained with acid fuchsin or light green in acidic solution and differentiated in water or weak alkali. When sufficient material was available, permanent microscopical preparations were made of the female subgenital plate and of the elytra, which were cleared as above and mounted on slides in PBS-glycerol solution. Measurements were made with a digital micrometer stage under 12X or 25X magnification. In the tables of dimensions, E is the length of the elytron, from the point at which the suture between the mesothoracic epimeron and episternum crosses the wing base to the tip of the elytron; F is the maximum length of the femur; P is the length of the pronotum in the dorsal midline, measured parallel to the line joining the highest points of the pronotum. We estimated sexual dimorphism by calculating the ratios of the male femur to the female femur, and so on for elytron and pronotum, finally averaging the three measurements.
Results
Morphology of the internal genitalia.—
Male.
The first descriptions of eumastacoid internal male genitalia were provided by Chopard (1920) and Grassé & Hollande (1946). Rehn's (1948) systematic review of the Eumastacoidea ignored these structures. Rehn & Grant (1958) made the first systematic treatment of the male genitalia of the entire Eumastacidae (s.s.) on the basis of which they erected three new subfamilies of the Eumastacidae (Paramastacinae, Pseudomastacinae and Temnomastacinae) and restricted the previously erected Eumastacinae. The only revision of the genus Eumastax (which then included Homeomastax) is that of Descamps (1971) and his is the only modern general systematic account of the whole superfamily (Descamps 1973a). For his description, Descamps uses anatomical terms in part deriving from Grassé & Hollande (1946) and Rehn & Grant (1958), many of which imply homology of at least some eumastacine and acridoid structures. We prefer to use more neutral terms for some of the major sclerites. In the following account we use “ectophallic sclerites” for Rehn & Grant's “ectophallic valves”, noting however (as pointed out by those authors) that they are situated at the junction of the endo- and ectophallic membranes, and it is not obvious to which they belong. We use “endophallic plate” for Descamps' (1971) “penis”; Descamps (1973)9 states (and we concur) that in the Eumastacidae the endophallus cannot be everted and does not form an intromittent organ, so “penis” seems inappropriate. We retain the terms epiphallus and epiphallic lophi, and spermatophore sac and ejaculatory sac, as even if the structures are not necessarily homologous with the acridoid structures of the same names, their function appears to be identical.
The general structure of the phallic complex of Homeomastax is illustrated in Fig. 4. It consists in essence of a short cylinder of membrane, the ectophallic membrane, closed at the posterior end. In the retracted condition, the membrane is reflexed outwards and backwards at the anterior end of the cylinder and inserts around the rim of the subgenital plate (Fig. 4B); in Homeomastax this membrane does not fold dorsally to form a visible pallium covering the aperture of the subgenital plate, though it apparently does so in some other eumastacine genera (Descamps 19739). The point at which the membrane folds ventrally is determined by a variety of stiffening structures in the different species; a medial longitudinal ridge of membrane (Fig. 4B), or an area of light sclerification of various extent and shapes (e.g., Figs 19D, 20A, 21B). There is often a change in membrane consistency at this point and Descamps (1971) refers to this division as the “repli ventral” or ventral fold (Fig. 4A). Internally, more or less axially within the outer cylinder, is the endophallic plate, a single sclerite, unitary anteriorly but dividing into two arms posteriorly, which bears the membranous sacs used for delivery of sperm (Figs 4C, F, G). The most distal sac, the spermatophore sac, opens dorsoapically through the ectophallic membrane along a slit, the genital aperture, which lies in a trough in the ectophallic membrane, the genital trough (the “fente phallotremique” of French authors) (Fig. 4E). Along the margins of the genital trough lie the paired ectophallic sclerites (Fig. 4C). These sclerites typically have the form of a bar, projecting posteriorly somewhat beyond the end of the genital trough. Laterally and apically the ectophallic sclerites extend as progressively thin sclerotized sheets, covering the posterior end of the phallus; we refer to these sheets as the apical shields (Figs 4E, F), and they typically bear basiconic sensilla and sometimes cuticular ornament. The posterior ends of the arms of the endophallic plate insert on the lower margins of the apical shields, close to the midline (Figs 4C, F). The insertion is flexible and not much sclerotized (Fig. 4G), and can be dissolved completely by over-vigorous maceration in alkali. This fact has given rise to erroneous descriptions of the arms of the endophallic plate in some species as being unusually short: in stained specimens which have been only lightly treated in alkali it can be seen that the endophallic plate always extends to the apical shield.
In the midline the ectophallic membrane forms a complex series of invaginations and evaginations. One of these invaginations is the genital trough (and possibly the spermatophore sac too, though we have no data about the ontogeny of this structure). The genital trough in many species is prolonged anteriorly, forming a much-pleated sac around the proximal ends of the ectophallic sclerites, the sclerite sac (Fig. 4B). Above this in most species the membrane forms a hollow, posteriorly-directed flap, the transverse dorsal fold (Rehn & Grant 1958), which partially covers the genital trough (Figs 4A, D); in some species the upper membrane of the dorsal fold is partially sclerotized. Above this there is usually a pocket-shaped invagination, the subepiphallic sac (Figs 4B, C) and above this again, the roughly oblong epiphallus, which is provided with two very large hook-shaped lophi (Fig. 4A). The epiphallus itself is apparently also an invagination of the ectophallic membrane, only lightly sclerified, and its posterior end hangs free in the haemolymph, with only the lophi projecting free into the cavity enclosed by the subgenital plate. It is thus probably not homologous, at least not in its entirety, with the similarly named acridoid structure, which lies completely in the plane of the membrane. (For this reason we are reluctant to use Eades' (2000) term “pre-epiphallic diverticulum” for any of the sacs we describe above). From the dorsal posterior margin of the epiphallus, the ectophallic membrane runs posteriorly and dorsally and is inserted at the base of the paraprocts. The epiphallus is highly flexible and rather variable in shape between individuals (see e.g., Figs 4H–K), though the shape of the lophi and of their basal sensory plaques have taxonomic value at the specific level.
Below the posterior end of the genital trough the ectophallic membrane is evaginated in the midline in many species to form a more or less elaborate ventroapical evagination (Descamps 1971) (e.g., Figs 10 C, E, G). This appears to be the structure named “ventral fold” by Rehn & Grant (1958); it is however not a fold, so Descamps' name appears more suitable. Note that Descamps (1971) sometimes also uses the term “repli ventral”, but in a different sense (see above). The dorsal proximal part of this structure is sometimes more or less continuous with the floor of the genital trough and so with that of the spermatophore sac. It can be a simple membranous sac (e.g., Figs 16C, H) or an elaborately shaped structure provided with cuticular ornamentation (Fig. 20D). In those species without a ventroapical evagination there are longitudinal pleats in the membrane at this point (e.g., Fig. 4E), which allow the ectophallic sclerites to separate and the genital trough to widen during copulation (see below). In most species, and in all with a large ventroapical evagination, a large Y-shaped apodeme inserts medially on the membrane just anterior to the lower edge of the evagination; its arms terminate on either side of the ejaculatory sac, below the endophallic plate (e.g., Fig. 9E).
During copulation the whole membranous complex is advanced towards the aperture of the cup-shaped subgenital plate, apparently by haemolymph pressure. As in other Caelifera, the male supra-anal plate or epiproct is depressed in copulation and then fits behind the posterior edge of the epiphallus. The inner proximal surface of the hooked lophus of the epiphallus forms a transparent dome-like plaque provided with numerous sensilla (see e.g., Figs 4D, 6E & F), which presumably could detect the presence of the epiproct. (The epiproct itself is not a rigid structure as in the Acridoidea, but is a soft membranous bag with numerous pleats and folds, the cavity of which is part of the haemocoel; it tends to shrivel into arbitrary shapes of no taxonomic value in dried specimens). When the epiproct is engaged with the epiphallus (and presumably stiffened by blood pressure) it must assist in keeping the genital mass advanced. The endophallus receives dorsally and proximally the ejaculatory duct leading from the testis, which communicates with two further sacs, named by Rehn & Grant (in conformity with acridoid practice) the proximal ejaculatory sac and the distal spermatophore sac (Fig. 4C). The former is accessed via the slot through the endophallic plate, and hangs ventrally below the plate, inclined towards the rear in most species. The spermatophore sac lies dorsally on the endophallic plate, and its floor lies between the arms of the latter; it is likely that the entire sclerite represents a sclerotization of the membrane of the sac. The posterior arms of the endophallic plate fuse with the apical shields, as described above, thus providing a skeletal framework around the posterior end of the spermatophore sac, which opens dorsally and posteriorly to the exterior by a slit at the bottom of the genital trough, between the ectophallic sclerites.
The musculature associated with the male genital apparatus is relatively simple and most of it seems to function only to retract the apparatus after its eversion. The main musculature is ventral and longitudinal and inserts on the Y-shaped apodeme which originates at the base of the ventroapical evagination; strands of this muscle also insert on the inside of the apical shields, especially in species lacking a medial apodeme. Additional muscles insert around the base of the hooked lophi of the epiphallus. No muscles insert on the endophallic plate, nor on the anterior parts of the epiphallus. The phallic musculature thus seems unlikely to function as a constriction mechanism to extrude the spermatophore precursors, as described for the Acridoidea by Snodgrass (1935) and Eades (2000). What may be glandular tissue occurs between the lateral walls of the spermatophore sac and the ectophallic membrane, and is richly supplied with tracheae.
Mode of operation of the male genitalia in copulation.—
In a specimen of H. robertsi we found a fully developed spermatophore lodged in the phallic complex (Figs 5D, 21F) The rigid, pointed and somewhat hooked tip of the spermatophore protruded from widely parted ectophallic sclerites and projected above the dorsal rim of the subgenital plate. The supra-anal plate was depressed and locked behind the epiphallus, as described above. After maceration in alkali the spermatophore was gone, but the phallic complex was left in its everted state (Figs 21G, H). Between the spread sclerites, the opening of the spermatophore sac is seen to form a circular aperture through which the spermatophore was extruded. The membranes below and posterior to the genital aperture are retracted in a series of folds, and the dorsal fold too is retracted from the genital slit. The ventroapical evagination, which in H. robertsi is highly elaborated in the shape of a lightly sclerotized spoon, covered with small denticles, is, however, still in the folded position in this specimen, and not extended. This, together with the depressed supra-anal plate suggests that this specimen was about to copulate, but not actually engaged, when killed.
We were also able to relax and dissect a pair of H. strigla n. sp. which had been killed in copulation and then pinned without separating the pair (Fig. 5). In copulation, the male abdomen is held in the normal position, apart from a slight axial rotation of the tip towards the female. The opening of the subgenital plate is appressed to the extruded aperture of the bursa copulatrix, which the female places on top of the male terminalia (see Fig. 5A–C, and next section). In the copulating pair the male supra-anal plate and phallic complex were as described in the previous paragraph, but additionally the ventroapical evagination was extended, forming a channel extending from the ventral rim of the opening of the spermatophore sac (Figs 5E, F). The tip of the evagination appeared to be lodged between the ventral ovipositor valves at their base. No spermatophore was evident in this case, suggesting that it had either not yet been formed, or had already been absorbed by the female.
The points of the hooked lophi project vertically upwards, and seem to be the only structures which could engage with the female genitalia to provide mechanical stability during copulation. However, in dissection of the pair killed in copulation we were unable to ascertain any contact between the lophi and the female. The hooked and pointed tip of the spermatophore, together with the absence of any other intromittent organ, suggest to us that the spermatophore itself might be used as a probe to enter the female reproductive tract, in the way that the aedeagal valves are used in the Acridoidea. This hypothesis would imply that the spermatophore would be formed within the male tract at the start of copulation; this view is supported by the specimen of robertsi described above, in which the fully formed spermatophore was almost entirely contained within the male spermatophore sac. It is, however, also possible that it is first formed within the female tract after copulation is under way, as in the Acridoidea, and that our observations on robertsi are an artifact of an interrupted copulation. Experiments on insects mating in captivity are required to resolve this question.
Morphology of the internal genitalia.—
Female. The female genitalia have been described by Descamps (1971, 1973a). The bursa copulatrix is completely membranous and heavily pleated, and occupies a space between the subgenital plate and the ventral valves of the ovipositor. It gives rise ventrally to a long, pleated medial structure, located between the lateral sclerified zones of the subgenital plate. Descamps (1971, 1973a) attributed to this structure a function similar to that of the “egg-guide” of the Acridoidea. Possibly this is indeed the case, but we note that it differs very considerably in morphology from the egg-guide, as it is simply a membranous fold, very delicate and without definite shape.
Dorsally the bursa copulatrix has a homogenous appearance, with lateral longitudinal folds and a small central zone where the folds take on a concentric form. This was found in all the individuals dissected. The form of these pleats suggests that at some physiological stage (probably during copulation) this area can be dilated. It gives rise to a duct of somewhat less delicate consistency, of constant bore and variable length, ending in an enormous amorphous ampulla. In all the species examined, the duct is shorter than the total length of the terminal ampulla, which in turn is very thin and delicate and without definite shape.
Descamps (1971) describes how in H. dereixi there is a small basal diverticulum and in H. bouvieri there is none. In the specimens of these (and other) species we examined there was no such diverticulum to be seen. Descamps stresses the extreme fragility of the structure and the reasons why the basal diverticulum may be inconspicuous. While allowing this, we do not agree that it is collapsed by the pressure of the enlarged oviduct in mature individuals, as he suggests. In all the cases we have examined, the ampulla is separated from the oviduct and adheres instead to the end of the alimentary tract. In fact, the large terminal abdominal ganglion of the ventral nerve cord is interposed between the terminal ampulla and the oviduct.
Mode of operation of the female genitalia during copulation.—
In the copulating pair mentioned in the previous section the tip of the female abdomen had been brought up alongside and slightly above the male genitalia, and then placed on top of the male apparatus. The complex of ovipositor valves and bursa was rotated axially through nearly 90° towards the male, effectively removing the female subgenital plate as a barrier to copulation. The opening of the bursa is thus exposed and additionally extruded, and it is this that is placed in contact with the male aperture, mediated by a torsion of the female abdomen (Fig. 5A–C). We imagine that the female senses the protruding tip of the spermatophore and inserts it into her orifice. Such behavior, together with the absence of interlocking mechanical structures (see above), suggest a large role for female choice.
Taxonomic significance of the female reproductive apparatus.—
In view of the fact that the spermatheca has no very definite shape, and that we find no appreciable differences in it between different species of Homeomastax, we consider it to be of little use as a taxonomic character, at least at the species level. In the following descriptions it is ignored. Similarly, we do not find systematic differences in the shape or armature of the ovipositor valves other than a smaller number of teeth on the dorsal valve in some smaller members of the surda species-group. The exact form and number of the teeth is variable, and often different on the left and right sides of the same individual.
Taxonomic value of other characters.—
Although Descamps (1971, 1973b) has used venation characters of the elytron and the shape of the pronotum in his descriptions, in most species we could not convince ourselves that the interspecific differences were larger than the intraspecific variation. We can detect differences in the prominence of the various ridges and carinae on the face, which seem to be species-specific, but these are too subtle to be readily described or used taxonomically. The number and relative sizes of the inner and outer spines of the hind tibia is also variable within species and to some extent between the left and right legs of individuals; although there are statistical differences between species, these characters rarely assist in identifying individual specimens. The coloration of the frons and basal antennal segments of both sexes varies in a species-specific manner in pinned dried specimens, and is given in our account; however, the relation of these colors to those in life is dubious. In most species the head and face are predominantly metallic blue in life, but this color fades soon after death. The colors in pinned specimens, other than the vermilion markings on the face which occur in females of some species, may be largely artifactual.
Species identification.—
Because there are so few useful external characters, we do not give a key to species. For species identification we recommend the use of Figs 2 and 3 (summary of male and female subgenital plates) and Table 23Table 23. (summary of dimensions and morphometric ratios), in which the ratio E/F is the most useful. For certainty, examination of the male phallic complex is usually essential.
Homeomastax Descamps 1979
Descamps 1979: 136.
Type species.— Eumastax surda Burr 1899.
Descamps 1971: 133, 137 (as the “surda group” of Eumastax).
Diagnosis.— A formal generic characterization was not given by Descamps (1979), but the key to genera he provided and his publication of 1971 indicate that his diagnostic characters were the following:
— Male antenna with 14 segments, female with 13–14; antennal organ on 7th segment from tip (differentiates from Araguamastax, Andeomastax, Santanderia, Erythromastax, Beomastax & Caenomastax);
— Macropterous (differentiates from Phrygonomastax and Hysteromastax);
— Male supra-anal plate subtriangular, without lateroapical angles (differentiates from Sciaphilomastax);
— No projecting pallium; femora distally red, or ringed red and black (differentiates from Eumastax);
— Ratio of length of elytron to length of hind femur (E/F) 0.5–1.0.
— Posterior margin of pronotum never notched medially; lateral carinae of metazona of pronotum subobsolete, not bordered ventrally by a black line (differentiates from Helicomastax Rowell & Bentos-Pereira, 2001).
Description.— (see also Fig. 1, habitus) Small-medium grasshoppers, body length 11.0–19.5 mm; F = 9.2–12.4 mm (males), 12.7–15.8 mm (females). Brachypterous to fully winged, mean E/F = 0.53–1.08. Sexual dimorphism in size (males/females) averages 0.79–0.89; ratio E/F always 4–15% larger in males: see Table 23Table 23..
Antennae with 11–12 flagellar segments, laterally compressed, black; scape and pedicel large, cylindrical, variously colored but not black. Eyes globular, protruding, almost hemispherical, in life strikingly colored, with the upper half black and the lower half white. Head approximately 3X as long dorsoventrally as anterioposteriorly; frons receding, straight in profile. Fastigium more or less absent, frontal ridge narrow, medially grooved between eyes, narrowest dorsal to medial ocellus, continuing to supraclypeal triangle, more or less indistinct ventrally. Infraorbital carinae well developed.
Pronotum more or less straight along anterior margin, posterior margin convex, smoothly rounded, never angular, truncate or medially notched. Medial carina very weak, especially in males; lateral carinae indistinct or obsolete, best developed in metazona. Lateral carinae cut by a single transverse sulcus, convex towards the rear in the pronotal disc, where, however, it is obsolete or reversed in profile. Dorsal profile of pronotum more or less flat (most males) or more concave and saddle-shaped (most females). Entire pronotum surrounded by a narrow thickened rim. Prothoracic episternum subquadrate in lateral view. Membrane of pro/mesopleuron bears a short lateral spine, weakly sclerotized, normally hidden behind posterior ventral angle of pronotum. Elytron diverging towards apex, tip rounded, usually suffused blackish at the base.
Legs long and slender. Fore and middle femora rounded on ventral face, square in profile on dorsal face, with two low longitudinal carinae terminating distally in short spines; anterior lobe of knee elongate and rounded, posterior lobe short and truncate. Fore and midtibiae quadrangular in section, with 10–12 internal and external spines. First tarsal segment almost as long as third segment, second segment very short; pulvillus large, tarsal claws well developed, with minutely serrate ventral edges. Hind femur with well developed outer, inner and medial carinae on both dorsal and ventral surfaces; all carinae except the ventral medial one are minutely denticulate. The dorsal and ventral carinae of the inner face join some distance before the knee to form a single carina. Brunner's organ present. Pregenicular region very slender; knee provided with 5 sharp spines, corresponding to the ends of the three most dorsal carinae and the ends of the ventral lobes of the knee. Tibia quadrangular in cross section, the upper edges densely haired, the lower provided with sharp spines extending from the knee to the tip; external and internal terminal spines present. External spines 14–26, all small and subequal in size; 18–24 internal spines, small proximally, large at midlength, then medium distally; the largest 5–6 spines alternating with smaller ones. On average (all species) 1–2 more outer spines than inner spines. Tarsal spurs present, straight, ventral edges minutely serrate; the external pair subequal, the more medial of the internal pair about 3X longer than the other three. Second tarsal segment very short, only about 10% of total length of foot; first tarsal segment longer than third, quadrangular in cross section, the upper edges terminating distally in two short spines, the inner extending somewhat slightly more posteriorly than the outer.
Male furcula absent. Male supra-anal plate membranous, complexly pleated and folded. Male cerci usually simple, slender, tapering, straight or slightly inwardly curved, rounded or spatulate at tips, in one species (H. cerciata) thick and obtusely angled downwards at half length. Paraprocts large, blackish, the extremities held more or less vertically, not lying horizontally under the supra-anal plate. Male subgenital plate simple, cup-shaped, rounded or pointed at posterior tip in dorsal view, often carinate in midline, entire to deeply notched at midline margin, varying between species (Fig. 2).
Female cerci and supra-anal plate similar to those of male, but the latter deeply sulcate medially and proximally. Ovipositor valves long, thin, pointed and prominently toothed. Dorsal valves widely separated at base, converging distally; dorsal face with rows of teeth, the largest along the outer edge; 10–13 teeth in outer row, including the apical tooth, exceptionally (H. carrikeri) 7–10; a large medial tooth opposite tooth 4 of the outer row (counting from the tip). Lower ovipositor valve with apical tooth strongly hooked; at least 4 major teeth along outer edge of distal portion of valve, but the proximal 3 often subdivided to give up to 7 teeth in total; a single prominent tooth on inner edge of distal portion of valve, pointed or bicuspid; typically 4–8 small teeth along both outer and inner edges of proximal portion of valve. Female subgenital plate very diverse, from simple triangular to trilobed and complexly spined, varying from species to species (Fig. 3). Most species have the lateral lobes of the distal extremity of the subgenital plate more or less thickened and sclerotized; this is best seen in cleared and mounted preparations, but is frequently visible externally as paired, darker areas of more rugose cuticle.
Coloration.—
When alive most species fit the following description, made from live H. robertsi; the different species vary in the color of the frons and genae, the stipe and pedicel of the antennae, and of the labrum and clypeus.
Male: general color, greenish yellow. Antennal flagellum, black. Upper half of eye black, lower half white. Head, light blue marked with black and metallic dark blue spots; V-mark on occiput, black; medial line on vertex and occiput green. Elytra and wings transparent, suffused blackish basally. Underside of metathorax and abdomen, subgenital and supra-anal plates, light blue. Cerci, black. Metallic yellow stripe laterally from mesonotum to 6th abdominal segment. Femora green, pregenicular area vermilion or red, sometimes diffusely ringed with a browner hue, hind knees brown or black. Hind tibia greyish, with a lighter ring below the knee.
Female: generally as male, but yellow of lateral stripe weaker, and blue and red colors duller, especially on head, where dorsal blue areas partially replaced by green. Black upper half of eye contains a white medial spot. Frons and basal antennal segments often marked or suffused with vermilion.
Coloration fades and changes markedly soon after death: most blue is lost, some of it turning grey or black, the entire eye turns brown and green areas become yellow.
Species descriptions.—
It is usual in revisions of genera to deal with the previously described species first and the new ones afterwards. We depart from this practice because the species of Homeomastax fall readily into species-groups, defined on the basis of the structure of the male phallic complex. We therefore deal with each of these groups in turn, listing first the old and then the new species for each.
The resultant order is the following:
A. surda species-group.
1. Homeomastax surda (Burr).
2. H. carrikeri (Hebard).
3. H. veraguae (Hebard).
4. H. bustum n. sp.
5. H. pluvialis n. sp.
6. H. acrita n. sp.
7. H. quinteroi n. sp.
B. bouvieri species-group.
8. H. bouvieri (Bolívar).
9. H. dereixi (Descamps).
10. H. jeanninae (Descamps).
11. H. brachyptera n. sp.
C. kressi species-group.
12. H. kressi (Rehn & Rehn).
13. H. hondurensis (Rehn & Rehn).
14. H. cerciata (Hebard).
15. H. tenoriensis n. sp.
D. silvicola species-group.
16. H. silvicola (Rehn & Rehn).
17. H robertsi (Descamps).
18. H. strigla n. sp.
E. Species known only from females.
19. H. dentata (Saussure).
20. H. richteri (Descamps).
21. H. annulipes (Descamps).
22. Homeomastax sp. indet.
The surda species-group (sensu novo, not in sense of either Rehn & Grant (1958) or Descamps (1971)).
Diagnosis.— Members of this group share the following characteristics:
— Phallus small.
— Ventroapical evagination small and simple, or even (H. surda) almost absent.
— Dorsal fold forming a unitary flap over the genital trough. Dorsal surface of this flap may contain paired sclerites (absent in surda, pluvialis n. sp. and bustum n. sp.).
— Sclerotized part of endophallic plate short and rather wide, rounded or angular at the anterior end.
— Epiphallus separated from dorsal fold by a relatively large area of membrane; subepiphallic sac present.
— Usually small in overall size, includes most of the smaller species of the genus.
— Pronounced sexual dimorphism in size (except in H. pluvialis n. sp.).
Homeomastax surda (Burr 1899)
Eumastax surdus Burr 1899: 90, 258, 264.
Type locality.— Chiriquí, Panama (nec. “Peru”) (holotype female).
Location of type specimen.— holotype: Naturhistorisches Museum, Vienna (Bruner collection).
Bruner 1901: 22, 23, pl. 2, Figs 14, 14a - as Eumastax surda, correction of gender.
Kirby 1910: 77.
Hebard 1924a: 92 (in error, misidentification of H. veraguae).
Hebard 1933: 126.
Rehn & Rehn 1934: 64 (redescription and allocation of male).
Descamps 1971: 139.
Descamps 1973b: 955.
Descamps 1979: 136. Transfers to Homeomastax; designates as type species of genus.
Rehn & Rehn (1934) wrote that in their opinion it would be impossible to determine the specific identity of the female holotypes of either H. surda (Burr) or H. dentata (Saussure), and did not examine them. Instead, they selected putative males of both species on the basis of either geographical proximity (H. dentata) or ecological similarity (H. surda) to the original type localities, and on this slender basis redescribed both species, though they refrained from designating their males “neallotypes”. Modern usage of these specific names has been based on the figures given by Rehn & Rehn (1934) and later by Descamps (1971), who also used the Philadelphia material of the former authors.
Examination of the holotype of H. surda shows that it is indeed very probably the same species as the population collected by Rehn in Pozo Azul de Pirrís, from which he selected his male. The reasons for thinking this are the following:
1. The subgenital plate. Both the holotype and the females from Pozo Azul have a somewhat elongated, sharply pointed subgenital plate, the margin of which is decorated with a few vertical denticles.
2. Dimensions and morphometric ratios (see Table 1). Comparison of the holotype with the sample of females from Pozo Azul, shows the former to be a small specimen with a relatively large pronotum, causing the morphometric ratios having P as a denominator to be smaller; in the case of E/P, significantly so. The E/F ratio of the holotype is equal to that of the Pozo Azul average. H. surda is found over a rather large geographical area, however, and seems to vary somewhat in size over this range. When the holotype is compared with a sample of females drawn from the entire Costa Rican range (from Surubres to the Panamanian border at Montezuma), it is found that none of the values for the holotype differs significantly from these means. Note that while the variance of the measured lengths increases in the larger sample relative to that from Pozo Azul, that of the ratios remains the same.
So far as is currently known, the only other species of Homeomastax likely to occur in Chiriquí (the type locality of H. surda) is H. acrita n. sp. It is highly improbable that the holotype of surda and H. acrita are the same taxon. In H. acrita the female subgenital plate is shorter than in H. surda and lacks marginal denticles. The E/F ratio for female H. acrita is 0.74 ± 0.03, 3 standard deviations away from the value for the holotype of H. surda.
Diagnosis.— Subgenital plates of males and females as in Figs 2 and 3. Female subgenital plate acutely pointed apically, lateral lobes well sclerified, separated medially by a depression; margins with a few small vertical spines, not visible in ventral aspect. Male subgenital plate in lateral view pointed, dorsal edge straight, margin not notched medially, in axial view with a short medial carina. Costal vein of elytron meets anterior margin more distally (at about 45% wing length) than in other species (about 25%).
Male internal genitalia as in Fig. 4. Epiphallus approximately square in dorsal view, with thin pointed lophi; posterior margin straight between the rather wide sensory plaques; anterior margin with a short truncated V-shaped membranous area. Subepiphallic sac present. Dorsal fold unitary, large, broad and square, posterior margin slightly convex. Sclerite sac present, large, up to twice as long as subepiphallic sac. Ectophallic sclerites as in Fig. 4C–E, resembling the profile of a bird's head in lateral view; pointed, slightly diverging at the tips in dorsal view. Spermatophore sac inserts at extreme anterior end of ectophallic sclerites. Ventroapical evagination minute or obsolete, its apodeme absent. Anterior end of endophallic plate rounded, narrowly winged (Fig. 4F, G). Ejaculatory sac relatively large (Fig. 4C).
Description.— Small in size, especially in males; only H. acrita n. sp. and H. brachyptera n. sp. have a shorter male femur on average (Table 1). Moderately long winged. Markedly dimorphic in size; male/female ratios 0.76–0.82, mean 0.79 (Table 23)Table 23.. Outer hind tibial spines, 20–21, inner spines, 20–22. Male frons whitish in dried specimens, basal antennal segments blue, in females yellow-orange; clypeus and labrum blue. Distal part of femur a clear red with little or no dark shadowing.
Distribution.— (Fig. 23) Lowland (up to 600 m) Pacific slope of Costa Rica, south of 10°N, and of extreme south western Panama, west of 82°W. There is little suitable habitat remaining in the Panamanian portion of this range. In Costa Rica however the range includes at least three National Parks, those of Carara, Quépos and Corcovado.
Material examined.— COSTA RICA
Puntarenas: Agua Buena: Campo Dos y Medio: Finca Hidalgo, 600 m, LS 295800_582000, 26 Aug 1995 (Rowell CHF Meier V), 1 ♀, 3 ♂ ♂, specimen nos 95377 95657_95659 (RC).
Puntarenas: Between Golfito and Gamba, along roadside through forest. LS 292500_551200, 30 Aug 1995 (Rowell CHF Meier V), 1 ♂, 3 ♀ ♀, specimen nos 95427 95428 95655 95656 (RC).
Puntarenas: between Villa Briceño and S. Miguel, 300 m, LS 300000_555200, 2 Sep 1995 (Meier V), 1♂, 1 ♀, specimen nos 95456, 95457 (RC).
Puntarenas: Golfito: fila above the city. 350 m, LS 288500_557500, 1 Sep 1995 (Rowell CHF Meier V), 2 ♂ ♂, specimen nos 95446 95654 (RC).
Puntarenas: Golfito: Fila Gamba, LS 291000_551000, 6 Sep 1979 (Rowell-Rahier M), 1♀, specimen no. 79013 (RC).
Puntarenas: Osa Península: Albergue Cerro de Oro, 270 m, LS 280000_518500, 16 March 1995 (Madrigal M), specimen no. CRI002 253146 (INBio).
Puntarenas: Osa Península: Est. Esquinas, 200m, LS 301400_542200, 15 May 1993 (Quesada JF), specimen no. CRI001 177293 (INBio).
Puntarenas: Osa Península: Fila Matajambre, LS 276600_525900, 7 May 1994 (Zumbado M), specimen no. CRI001 964324 (INBio).
Puntarenas: Osa Península: Llorona, LS 282000_498650, 18 Jan 1978 (Rowell CHF), 1 ♂, specimen no. 78043 (RC).
Puntarenas: Osa Península: Rancho Quemado, 200m, LS 292500_511000, 15 July 1992 (Quesada JF), specimen no. CRI000 736102 (INBio).
Puntarenas: Osa Península: Rincón de Osa, LS 294100_517800, 23 Jan 1973 (Rowell CHF) (RC).
Puntarenas: Osa Península: Rincón de Osa, 22 Feb 1966 (Roberts HR), 1 ♂, 4 ♀ ♀ (ANSP).
As above, but 18 Feb 1965, 1 ♀; 19 Feb 1965, 1 ♀.
Puntarenas: Osa Península: Rincón de Osa, 3–10 miles SW, arboreal, obtained by insecticide, 14 Feb 1967 (Roberts HR & EH, Harrison MS Moss WW Nickle D), 3 ♀ ♀ (ANSP).
Puntarenas: Osa Península: Rincón de Osa, 5 km. S, 8 Aug 1968 (Hespenheide H), 1 ♀ (ANSP).
Puntarenas: Osa Península: Sirena, LS 276600_513250, 20 Mar 1995 (Azofeita A), specimen no. CRI002 234059 (INBio).
Puntarenas: Osa Península: Aguas Buenas, 5 km. W. of Rincón de Osa, LS 295500_515500, 14 Sep 1995 (Rowell CHF Meier V), 3 ♂ ♂, 1 ♀, specimen nos 95556, 95557 95652, 95653 (RC).
Puntarenas: P.N. Carara: Estación Quebrada Bonita, 50 m, LN 195200_469400, 2 Aug 1991 (Rowell CHF Elsner N), 2 ♂ ♂, 2 ♀ ♀, specimen nos 91260_91263 (RC).
Puntarenas: P.N. Carara: Estación Quebrada Bonita. 50 m, LN 195200, 470000, 27 Sep 1993 (Rowell CHF), 2♂ ♂, specimen nos 93357, 93358 (RC).
Puntarenas: P.N. Carara: Sector Laguna Meandrica, 100 m, LN 197900_472800, 15 Jun 1990 (Zuñiga R), specimen no. CRI000 223733 (INBio).
Puntarenas: Quépos: P.N. Manuel Antonio, 80 m, LS 370900_448800, 15 Apr 1992 (Cano C), specimen no. CRI001 718654 (INBio).
Puntarenas: Uvita: above Finca Piñuela, 350 m, LS 342000_499000, 17 Sep 1995 (Rowell CHF Meier V), 1 ♀, specimen no. 95586 (RC).
Puntarenas: Villa Neilly: fila 2 km S. of Montezuma, 575 m, LS 291000_587000, 3 Sep 1995 (Rowell CHF Meier V), 1 ♂, 1♀, specimen nos 95476 95477 (RC).
San José: Pozo Azul de Pirrís, 325–550 ft (98–167 m) LS 400200_431200, May 1902 (Carriker MA), 3 ♀ ♀ (ANSP).
As above but 21 Aug 1927 (Lankester CH Rehn JAG), 5 ♂ ♂, 8 ♀ ♀, 6 larvae (ANSP).
San José: Pozo Azul de Pirrís, LS 400200_431200, no data (probably originally 21 Aug 1927, Lankester & Rehn) (MHNG).
San José: San Mateo: Río Surubres: Bonnefil Farm, 700 feet, LN 215500_476500, 17 Oct 1909 (Calvert PP), 1 ♀, (ANSP).
San José: San Mateo: Surubres, 250 m, LN 216200_477800, Jan 1906 (Biolley P), 1 ♀, 1 larva (ANSP).
PANAMA
Chiriquí: (Staudinger) (Vienna) (♀ holotype).
The specimen referred to by Bruner (1901: 23) from Panama: Chiriquí: Bugaba (coll. Champion) is no longer present in the Vienna collection.
Homeomastax carrikeri (Hebard 1923)
Eumastax carrikeri Hebard 1923: 181.
Type locality.— Colombia: Dept. Chocó: Andagoya, Antioquia, ♂ holotype and ♀ allotype. Types examined.
Location of type specimens: ANS Philadelphia.
Descamps 1971: 138, 155.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate with a prominent medial carina proximally, distally narrowed and produced into a long tapering tip. Male subgenital plate without medial carina, only slightly notched medially, subquadrate in axial view, dorsal edge convex in lateral view. Male cerci short and straight, slightly thickened at the tip.
Internal genitalia (Fig. 6) most similar to those of H. surda, but much less sclerotized. Epiphallus (Fig. 6E) more or less oblong in dorsal view, with a membranous central area; lophi thick and blunt, only shallowly hooked. Subepiphallic sac absent. Dorsal fold (Figs 6A, D) simple, short, rather thick in lateral view, without sclerites. Sclerite sac well developed. Ectophallic sclerites only lightly sclerotized, confined to upper edge of genital trough, of simple bar-like shape, diverging slightly but not projecting posteriorly. Apical shields minimal, simple. Ventroapical evagination (Fig. 6C) very small, but provided with a medial apodeme. Endophallic plate (Fig. 6G) very wide, rounded in shape at its anterior end, almost circular, the wide and smooth flanges extending far down the arms. Ejaculatory sac unusually large, spermatophore sac small.
Description.— The smallest species of the genus, relatively long winged, especially in males (Table 2). Sexual dimorphism in size 0.80–0.93, mean 0.88 (Table 23)Table 23.. Male with 20–21 outer tibial spines, 18 inner ones; female with 14–17 outer spines, 18–19 inner ones. Face white in dried specimen, genae blackish brown; scape, pedicel, labrum and clypeus were probably blue. The red part of the hind femur shadowed by two imperfect brown rings.
Distribution.— (Fig. 24) Colombia: Southern Chocó.
Homeomastax veraguae (Hebard 1933)
Eumastax veraguae Hebard 1933: 127.
Type locality.— of holotype ♂, Panama: Colón: “Guamina” (probably in error for Juan Mina, 9°09′N, 79°40′W), Río Chagres; of neallotype ♀, Panama: Colón: Porto Bello (= Portobelo).
Location of type specimens.— ANS Philadelphia (holotype lost).
Hebard 1923; 178 as Eumastax cerciata, partim (misidentification).
Hebard 1924; 92 as Eumastax surdus, partim (misidentification).
Rehn & Rehn 1934: 68; designation of neallotype female.
Descamps 1971: 138.
Descamps 1979: 136: transfers to Homeomastax.
The holotype male was originally identified by Hebard as H. surda (1924) and only later (1933) recognized by him as a separate species. Rehn & Rehn (1934) described as the female, a specimen from the type series of H. cerciata Hebard 1923. The conspecificity of the holotype and this female was once again only speculative, based on geographical proximity and similarity of both sexes to the Costa Rican surda. The holotype male was not available for our examination. It is currently lost; it may eventually resurface in either Philadelphia or Paris, its last certain location, and we refrain here from designating a neotype. We have examined the female described by Rehn & Rehn (1934) and a nonparatypic male from Barro Colorado Island, examined by these authors and referred by them to this species in the same paper. As they noted, the tip of the abdomen of this specimen has been crushed laterally, and the original shape of the subgenital plate is not apparent. It was however possible to extract the phallic complex. This is highly characteristic and agrees with fresh material recently collected from the former Canal Zone, including that of males caught in copulation. Of these latter, the male fits both the original descriptions and Descamps' (1971) drawings of the subgenital plate of the lost holotype, and the phallic complex is identical with that of the BCI specimen. The female of the pair is the same species as Rehn & Rehn's specimen; their speculation as to its identity is thus confirmed as being correct.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate with lateral lobes only barely sclerified, rounded at tip, with a slight medial carina, almost no marginal teeth. Male subgenital plate almost identical with that of H. surda, but with a slight medial notch, a more pronounced carina, and slightly more convex dorsal edge; cerci slightly longer.
Male internal genitalia (Fig. 7) most similar to those of H. acrita n. sp., but lacking the ornamented apical shields of that species. Epiphallus oblong, with roughly parallel, slightly convex sides. Lophi positioned more horizontally than usual in the genus. Subepiphallic sac of variable development, but usually present. Dorsal fold simple, with two long thin sclerified plaques on the dorsal surface (Fig. 7G). Sclerite sac poorly developed. Ectophallic sclerites obliquely and sinuously truncate in lateral view (Fig. 7D), strongly curved in dorsal view, with both posterior and anterior ends pointing inwards (Fig. 7G). Ventroapical evagination small, present; a deep ventral medial invagination lies behind it (Fig. 3 D, F, H). Endophallic plate either oblong (Fig. 3E) or rounded in shape at its anterior end, the flanges extending far down the arms.
Description.— Of medium size for the group (Table 3). The longest winged species of the surda species-group. Markedly sexually dimorphic in size (male/female ratios 0.76–0.83, mean 0.80 (Table 23)Table 23.. Hind tibial spines (female), 23 outer and 21 inner, 26 and 22 in a male. Face white tinged blue in dried male specimens, white with vermilion-tinged infra-orbital carinae in females. Red part of femur shadowed with brown distally and proximally, leaving only a narrow band of clear color.
Distribution.— (Fig. 23) Apparently confined to the lowland area in the provinces of Colón and Panamá corresponding roughly to the former Panama Canal Zone, and the region of Chagres National Park to its East, and including the islands of the Archipiélago de las Perlas. Although the specific name suggests an origin in the modern province of Veraguas, that term was used historically for all of Western Panama, including Portobelo (D. Quintero, pers. comm.), and it is presumably in this sense that Hebard used it. The species has not been recorded from Veraguas province.
Material examined.— PANAMA
Prov. Colón:
Porto Bello, 18 Apr 1912 (Busck A), 1 ♀(ANSP). This is the specimen described by Rehn & Rehn (1934).
Gamboa, Pipeline Road, 100–200 m, 4 Sep 1997 (Rowell CHF), 1 ♀, specimen no. 97270 (RC).
Gamboa, Pipeline Road, 15 Oct 97 (Rowell CHF, Bentos-Pereira A), 1 ♀, specimen no. 97358 (RC).
Plantation Road, between Summit and Gamboa, 12. Oct 97 (Rowell CHF), 1 ♀, specimen no. 97349 (RC).
Fort Sherman, 3 Sep 1999 (De Gracia L), pair in cop., specimen nos 99288, 99289 (UP). As above, but 27 Aug 1999, 2 pairs in cop., specimen nos 99548–99551 (UP).
Prov. Panamá:
Barro Colorado Island, 20 Jul 1924 (Banks N), 1♂ (MCZH).
As above but 24 Dec 1928 (Curran CH), 1 pair in cop. (ANSP).
As above but 14 Feb 1929 (Curran CH), 1 ♀ (ANSP).
As above but 7 Mar 1956 (Rettenmeyer), 1♀ (ANSP).
Archipiélago de las Perlas, Isla Contadora, 31 Dec 1972 (Engleman D), 1 ♂ (USNM).
Prov. Panamá: Altos de Cerro Azul, Los Nubes, 7 Oct 2000 (Quintero D), 1 pair in cop., specimen nos 2000187, 2000188 (UP).
Homeomastax bustum n. sp.
Holotype.— ♂: COSTA RICA: Puntarenas: Tárcoles: Reserva Biológica Carara, Sendero de los Enterrados, 450–530 m, 27 Jul 1980 (Rowell CHF & Rowell-Rahier M), specimen no. 80034 (ANSP).
Paratypes.— 1 ♂, 2 ♀ ♀, data as above, specimen nos 80033 80035 80036 (ANSP). As above, but Sendero Carara, 500 m, 26 Jul 1980, 1 ♂, 2 ♀ ♀, specimen nos 80030 to 80032 (RC).
Other material examined.— COSTA RICA: Puntarenas: Osa Península: Rincón de Osa, stream trail S. side of airstrip, 22 Feb 1966 (Roberts HR), one ♂, one ♀(ANSP).
Etymology.— Latin bustum, noun in apposition, place where the dead are burnt or buried, from the type locality Sendero de los Enterrados.
Diagnosis.— Can be distinguished with certainty only by the male internal genitalia. Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate almost identical with that of H. surda, but shorter, somewhat more heavily sclerotized at tip. Margin decorated with vertical spines, as in H. surda. Male subgenital plate not distinguishable from that of H. acrita, bluntly pointed, dorsal margin slightly convex in lateral view, medially notched and carinate.
Male internal genitalia as in Fig. 8. Ectophallic sclerites terminating posteriorly in long curved points (Figs 7C, D), obviously diverging in dorsal view (Fig. 8G). Subepiphallic sac present. Dorsal fold simple, truncate. Sclerite sac voluminous, longitudinally pleated, extending ventrally almost to posterior end of sclerite. Apical shields small, simple, not ornamented as in H. acrita. Endophallic plate (Fig. 8F) similar to that of H. acrita, oblong and slightly notched at anterior end, broadly winged. Ventroapical evagination small, somewhat pointed, with medial apodeme (Fig. 8E).
Description.— One of the larger of the surda species-group (Table 4). The specimens from the Osa are nearly 10% larger than those from Carara. Elytra of medium length; relative length the same as for H. surda, longer than in H. acrita, shorter than in H. veraguae. Sexual dimorphism in size well developed: male/female ratios 0.80–0.85, mean 0.82 (Table 23)Table 23.. Hind tibial spines (female), 24 outer and 21 inner, 26 and 22 in a male. Face and basal antennal segments yellowish white in the only nondiscolored male specimen, but may be faded; red portion of femur without rings or shading. Females all discolored.
Distribution.— (Fig. 23) To date known certainly only from the higher, inland part of the Carara biological reserve and from a single locality in the Osa Peninsula. The latter is the only known locality where two species of Homeomastax apparently occur sympatrically — other specimens collected at the same time and place are H. surda.
This is probably the species to which belong the specimens described erroneously as H. dentata (Saussure) by Rehn & Rehn (1934). The corresponding localities are not far from Carara, and their drawings of the male subgenital plate are compatible.
Homeomastax pluvialis n. sp.
Holotype.— ♂: COSTA RICA: Cartago: P.N. Tapantí, in forest along roadside 0–1 km past park entrance, 1500–1600 m, in cop., 8 Oct 1999 (Rowell CHF), specimen no. 99414 (ANSP).
Paratypes.— 1 ♀, data as above, in cop. with holotype, specimen no. 99415 (ANSP).
As above, but 20 Aug 1997 (Rowell CHF Singh I), 3 ♀ ♀, specimen nos 97094, 97095 (ANSP), 97096 (RC).
Etymology.— Latin pluvialis, of rain, referring to the exceedingly wet climate of the type locality at Tapantí.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate compressed laterally, appearing very narrow in ventral view, medially carinate proximally. All specimens examined have a heavily sclerotized, almost black tip; the paired thickenings of the lateral lobes are seen only in cleared preparations. Male subgenital plate most similar to that of H. surda; differs from latter in the somewhat more rounded tip in lateral view, the very short medial carina, and in having a black border along the dorsal edge. Cerci shorter and blunter than all other surda-group species apart from H. carrikeri.
Male internal genitalia as in Fig. 9. Ectophallic sclerites terminating posteriorly in a ventrally inclined projection in lateral view (Figs 9C, D), in dorsal view broadening at the tips, somewhat divergent, with a complex folded surface unlike that of any other species (Fig. 9G). Subepiphallic sac present (Figs 9C, D). Dorsal fold simple, somewhat pointed in the midline (Figs 9C, G). Sclerite sac present, extending ventrally about 40% of the distance to the posterior end of sclerite (Fig. 9D). Apical shields simple. Endophallic plate (Fig. 9F) somewhat rounded anteriorly, with a pleated lateral wing. Ventroapical evagination small, with medial apodeme (Fig. 9E).
Description.— One of the largest members of the surda species group, but with noticeably short wings; only H. brachyptera from Panama has relatively, still shorter ones (Table 5). Sexual dimorphism in size slight: male/female ratios 0.84–0.94, mean 0.89 (Table 23)Table 23.; note however that only a single male has been measured. Hind tibial spines (female), 24 outer and 22 inner, 21 and 21 in the male. Frons and basal antennal segments whitish in dried male, in females frons whitish, basal antennal segments blue with a vermilion spot at the joint, vermilion marks on infraorbital carinae. Red portion of femur shadowed with brown proximally and distally in both sexes.
Distribution.— (Fig. 23) To date known with certainty only from the type locality, in montane rain forest in the north-eastern Talamanca mountains of Central Costa Rica, on the Atlantic side of the watershed. A brachypterous Homeomastax species has been seen on the Pacific side near Cerro Pittier and may be this species.
Homeomastax acrita n. sp.
Holotype.— ♂: Puntarenas: S. Vito de Jaba, Finca Las Cruces, 1100 m, LS 304500_577500, 7 Aug 1980 (Rowell CHF & Rowell-Rahier M), specimen no. 80671.
Allotype.— ♀, in cop. with holotype, specimen no. 80672 (both ANSP).
Etymology.— Greek akritos, confused, mixed up, doubtful, an allusion to the taxonomic history of this taxon.
Rowell et al. 1983: 207 (ecology - as H. dentata, in error, following Rehn & Rehn (1934) and Descamps 1971).
Rowell 1998: (distribution - as H. dentata, in error, following Rehn & Rehn (1934) and Descamps 1971).
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate similar to but shorter than that of H. surda; distinguished from surda and bustum by absence of marginal spines. Male subgenital plate not distinguishable from that of H. bustum: relatively long, dorsal edge slightly convex in lateral view, slightly truncate at tip; small medial notch in axial view, thereby distinguished from H. surda.
Male internal genitalia as in Fig. 10. Epiphallus slightly longer than broad, outer margins convex, with an anterior membranous field; posterior margin almost straight between widely spaced sensory plaques, slightly pointed in midline (Fig. 10A). Dorsal fold simple, with two small sclerotized patches on dorsal surface (Fig. 10G). Ectophallic sclerites as in Figs 10C & D, obliquely truncate in lateral view. Spermatophore sac inserts about one third of the way along the ventral surface of the ectophallic sclerites (Fig. 10D). Highly distinctive rugose ornamentation on lateral portions of apical shields (Figs 10C, G). Ventroapical evagination small but present. Anterior end of endophallic plate (Fig. 10F) rounded laterally, flat or notched apically, broadly winged.
Description.— The smallest Central American species, only H. carrikeri (Colombia) is smaller (Table 6). Shorter winged than H. surda. Markedly dimorphic in size: male/female ratios 0.76–0.82, mean 0.79 (Table 23)Table 23.. Outer and inner hind tibial spines, 23 (female) or 20–21 (male). Male frons and basal antennal segments whitish in dried specimens, in females vermilion. Red part of femur shadowed with brown distally and proximally, leaving only a narrow band of clear color.
Distribution.— (Fig. 23) Highland (above 700 m) southern Pacific slope of Costa Rica, south of 9°20′ N, extending to the Panamanian border at Cañas Gordas and probably into Western Panama. All known localities are on the eastern sides of the basin of the complex of rivers that flows into the Pacific as the Rio Grande de Térraba, from the Valle de El General and the Valle de Coto Brus. The Fila Costeña bounds this basin to the west; on its seaward side H. surda is found.
Material examined.— Paratypes:
Puntarenas: S. Vito de Jaba, Finca Las Cruces, 1100 m, LS 304500_577500, 28 Aug 1976 (Rowell CHF), 1 ♂, specimen no. 76099 (MNHNP).
As above, but 15 Mar 1978 (Rowell CHF), 1 ♀, specimen no. 78098 (ANSP).
As above, but 31 Aug 1979 (Rowell CHF Rowell-Rahier M), 1 ♂, specimen 79009 (ANSP).
As above, but 27 Oct 1979 (Braker HE), 1 ♂, specimen 79012 (MNHNP).
As above, but 1 Aug 1980 (Rowell CHF Rowell-Rahier M Hyde C Braker HE), 3 ♀ ♀, specimen nos 80674, 80675, 80676 (MNHNP).
As above, but 1 Aug 1980 (Rowell CHF Rowell-Rahier M Hyde C Braker HE), 1 ♀, specimen no. 80677 (ANSP).
As above, but 3 Aug 1980 (Rowell CHF, Rowell-Rahier M, Hyde C, Braker HE), 1 ♂, specimen no. 80682 (MNHNP).
As above, but 5 Jun 2000 (Rowell CHF), 1 ♂, specimen no. 2000.032 (ANSP).
Puntarenas: San Vito: Fila Cruces, above R. Chico, 1430 m, LS 305900_569130, 9 Aug 1980 (Rowell CHF), 1♂, 1 ♀, specimen nos 80669, 80670 (ANSP).
Other material:
COSTA RICA
Puntarenas: Altamira, above Finca Colorado, 1450 m, LS 331400_571900, 4 Jun 2000 (Rowell CHF), 1 ♂, 1 ♀, 2 larvae, specimen nos 2000.021 to 2000.024 (RC).
Puntarenas: Cañas Gordas: Finca Loma Linda, 1075–1085 m, LS 299200_581100, 4 Aug 1980 (Rowell CHF, Rowell-Rahier M, Hyde C, Braker HE) (RC).
Puntarenas: Coto Brus: Est. Biol. Las Alturas, 1500 m, LS 322500_591300, 15 Apr 1992 (Araya F), specimen no. CRI000 574629 (INBio).
As above, but 15 Oct 1991 (Ramirez M), specimen nos CRI000 574629 & CRI001 209151 (INBio).
Puntarenas: Las Alturas, 1600 m, LS 323400_591300. 5 Sep 1995 (Rowell CHF Meier V), 1 ♀, specimen no. 95487 (RC).
Puntarenas: Las Alturas: lower slopes of Cerro Irkibi, 1500–1700 m, LS 323000_593000, 7 Aug 1980 (Rowell CHF Rowell-Rahier M), 1 ♀, 3 ♂ ♂, specimen nos 80686 to 80689 (RC).
Puntarenas: Las Alturas: trail above gravel pit, 1700 m, LS 324200_589200, 6 Sep 1995 (Rowell CHF Meier V), 4 ♂ ♂, specimen nos 95492, 95493, 95702, 95703 (RC).
Puntarenas: San Vito de Jaba, 6 km S, 08° 42′N, 83° 00′W, 21 Mar 1967 (OTS Adv. Zoo. Course), specimen no. CRI001 013406 (INBio).
Puntarenas: S. Vito de Jaba, Finca Las Cruces, 1100 m, LS 304500_577500, 28 Aug 1976 (Rowell CHF), 1 ♂, specimen no. 76100 (RC).
As above, but 27 Oct 1979 (Braker HE), 1 ♂, specimen no. 79010 (RC).
As above, but 31 Aug 1979 (Rowell CHF Rowell-Rahier M Braker HE), 1 ♂, 1 ♀, specimen nos 79011 (RC).
As above, but 1 Aug 1980, 6 ♀ ♀, specimen nos 80673 80679 (RC).
As above, but 2 Aug 1980, 1 ♀, specimen no. 80681 (RC, alcohol specimen).
As above, but 5 Jun 2000 (Rowell CHF), 1 ♂, specimen no. 2000.031 (RC).
Puntarenas: San Vito: Fila Cruces: Finca Ilana, above R. Claro, 1430–1280 m, LS 304500_570100, 25 Aug 1995 (Rowell CHF Meier V), 2 ♂ ♂, alcohol specimen nos 95700, 95701 (RC).
As above but 26 Aug 1995 (Chacón I), 2 ♂ ♂, 1 ♀, specimen nos 95383–95385 (RC).
Puntarenas: Sonador: Finca Longo Mai, 700 m, LS 357500_520500, 23 Aug 1995 (Rowell CHF Meier V), 1 ♀, specimen no. 5338 (RC).
S. José: Quisarrá: Finca Skutch, 695 m, LS 365300_505000, 29 Jul 1980 (Rowell CHF Rowell-Rahier M), 1 ♀, specimen no. 80683 (RC).
San José: Valle de El General: R. Unión: 3 km N. of Zapotal, 1345 m, LS 367600_517800, 7 Jun 2000 (Rowell CHF), 4 ♂ ♂, specimen nos 2000.057 to 2000.060 (RC).
Homeomastax quinteroi n. sp.
Holotype.—♂: PANAMA: Prov. Darién: P.N. Darién: Pirre: Estación Río Frio, 12 Nov 2000 (Cambra R Santos A), specimen no. 2000178 (ANSP).
Paratypes.— Same data as holotype, 2 ♂ ♂, 3 ♀ ♀, specimen nos 2000173, 2000178, 2000179, 2000182, 2000183, 2000186 (ANSP); specimen nos 2000170, 2000171, 2000173, 2000175, 2000176, 2000183; 7 ♀ ♀, specimen nos 2000172, 2000179, 2000181, 2000182, 2000184, 2000185, 2000186 (all UP).
Same data, but 29 Mar 2000 (Cambra R Santos A Bermúdez S), 1 ♂, 1 ♀, specimen nos. 2000168 & 200132 (UP); same data, 1 ♂, 1 ♀, specimen nos. 2000169 & 2000133 (RC).
Etymology.— Named for Dr. Diomedes Quintero, director of the Museo de Invertebrados G.B. Fairchild, Universidad de Panamá, in recognition of his collegial assistance of our field work in Panama and his valued help in obtaining specimens.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate distinctive, narrowing abruptly immediately anterior to lateral lobes, then running more or less parallel-sided to a bluntly rounded tip. Margin devoid of spines. Male cerci slender, pointed, slightly upwardly inflected in lateral view, more or less straight in dorsal view. Male subgenital plate of same type as in H. bustum and H. veraguae; short, dorsal edge in lateral view straight or very slightly convex; tip rounded in profile, then dropping in two straight sections to suture; in axial view upper margin slightly notched medially, with a long and strong medial carina running from the notch ventrally; in dorsal view tip bluntly rounded, the median carina projecting as a ridge.
Internal male genitalia as in Fig. 11. Epiphallus in dorsal aspect oblong or weakly trapezoid, tapering anteriorly; anterior margin with a deep membranous V, posterior margin straight; sensory plaques large. Subepiphallic sac present. Dorsal fold simple, but in some specimens medially notched, sometimes almost bilobed. Ectophallic sclerites simple, in form of vertical plates, in lateral view rounded at upper posterior angle, posterior edge vertical; in dorsal view, first converging then diverging apically. Apical shields small, oval, diffuse. Sclerite sac present, small. Ventroapical evagination small, projects posteriorly as a small pointed membranous lip below the ectophallic sclerites; medial apodeme weak, diffuse, not visible in all preparations. Endophallic plate flat, shallow, oblong at proximal end, with a wide, straight-flanged margin. Ejaculatory sac narrow, straight, vertical or inclined anteriorly; spermatophore sac large, longitudinally pleated.
Description.— A medium-large member of the surda group, long winged and long legged (Table 7). Markedly sexually dimorphic in size, male/female ratios = 0.77–0.81, mean 0.79 (Table 23)Table 23..
Distribution.— Known to date only from the type locality in Darién.
The bouvieri species-group
Diagnosis.— Members of this group share the following characteristics:
— Ventroapical evagination small or absent, medial apodeme absent.
— Epiphallus connected almost directly to base of dorsal fold, and appressed to it, covering it almost completely; subepiphallic sac usually short.
— Ectophallic membrane often with paired pigmented patches posteriorly and laterally.
— Apical shields small and simple.
— Dorsal fold without sclerotized plaques, forming a unitary flap over the genital trough.
— Sclerotized part of endophallic plate rounded at its anterior end, flange narrow.
— Apex of subgenital plate of females often trilobed (but not invariably— see H. dereixi).
The bouvieri-group resembles the surda-group in having some of the smallest and most brachypterous species of the genus and in the weak development of the ventroapical evagination. In its other characters it is clearly distinct, especially in the relatively immobile epiphallus, which is tightly appressed to the dorsal surface of the ectophallus. Most members of the group are S. American; only H. brachyptera n. sp. is Panamanian. H. richteri Descamps, the male of which is unknown, is probably a further member of this group, in view of its ornamented, trilobed, female subgenital plate.
Homeomastax bouvieri (Bolívar 1918)
Eumastax bouvieri C. Bolívar 1918: 22.
Type locality.— Ecuador: Santo Domingo de los Colorados, 510 m.
Location of type specimen.— MNHN Paris.
Descamps 1971: 138, 153.
Descamps 1973a: 216, Fig. 2.
Descamps 1973b: 948.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate long, trilobate at tip, central lobe sharply pointed. Male subgenital plate in side view similar to that of H. acrita, but in axial view distinguished by absence of median carina and very small medial notch.
Male internal genitalia as in Fig. 12. Epiphallus tapering anteriorly, somewhat emarginate on posterior margin, or with a shallow medial membranous area that stains more lightly than the bridge; tightly appressed to dorsal fold, which is simple and angulate in the midline (Fig. 12A). Ectophallic sclerites somewhat convex in dorsal view, bowed out laterally at midlength, terminating posteriorly in simple straight points, narrow in dorsal view, wider and blunter in lateral view (Figs 12B, C). Apical shields simple and small, decorated ventrolaterally with paired patches of black pigment (Fig. 12C). Endophallic plate rounded at anterior end, smoothly winged (Fig. 12C). Ventroapical evagination very small, almost absent, apodeme absent. The details of the junction of sclerite sac and spermatophore sac could not be determined without the risk of damaging the borrowed specimen. Subepiphallic sac, if present, very shallow.
Description.— A medium sized species with long wings (Table 8). Markedly sexually dimorphic in size: male/female ratios 0.68–0.78, mean 0.74, in the single Ecuadorian pair examined, but 0.78–0.84, mean 0.81, in Colombian specimens (Descamps 1971, see also Table 23Table 23.). The values of the Ecuadorian male measured in this study differ considerably from the average values given by Descamps (1971) for Colombian specimens, although the values for females are similar.
Distribution.— (Fig. 24) Ecuador and Colombia (Depts Nariño, El Valle).
Natural history.— On ferns (Descamps 1971).
Material examined.— Ecuador: Pichincha: P.V. Maldonado: Res. ENDESA, 700 m, 16 Jul 1991 (Amédégnato C, Poulain S), 1 ♂, 1 ♀ (MNHN, Paris).
Homeomastax dereixi (Descamps 1971)
Eumastax dereixi Descamps 1971: 139, 156.
Type locality.— Colombia: Dept. Córdoba: Ayapel, Caño Seco.
Location of type specimen.— MNHN Paris.
Descamps 1979: 136; transfers to Homeomastax.
Diagnosis.— Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate similar to that of H. surda, rather sharply pointed, not trilobate. Male subgenital plate in axial view essentially identical to that of H. bustum, in side view less elongate.
Internal male genitalia as in Fig. 13. Epiphallus with more or less straight posterior margin, not deeply emarginate. Subepiphallic sac small and shallow. Dorsal fold short and simple, rounded in dorsal view, almost completely covered by epiphallus. Ectophallic sclerites strongly divergent, ending posteriorly in a sharply pointed, downwardly curved tip (“hawk's beak”) in lateral view, as sharp divergent points in dorsal view (Figs 13A–C), deeply grooved longitudinally on their interior face (Fig. 13D). Apical shields simple, small, without lateral pigmented spot. Sclerite sac large, longitudinally pleated. Endophallic plate (Fig. 13A) rounded at anterior end, broader than in bouvieri or brachyptera n. sp. Ventroapical evagination absent.
Description.— One of the larger species of the genus, and with the longest wings, equivalent to those of H. veraguae (Table 9). Only moderately sexually dimorphic: male/female ratios 0.80–0.84 (Descamps 1971, Table 23Table 23.). Females have proportionately somewhat longer wings and femora.
Distribution.— (Fig. 24) Colombia: Depts. Córdoba, Magdalena, Cundinamarca (the latter awarded subspecific rank as H. d. cundinamarca by Descamps 1971).
Material examined.— COLOMBIA: Cordoba: between Monteria and Tierra Alta: Hda. El Torno de Matamoros, 1 Jan 1968 (Descamps M), paratype ♂ and ♀ (MNHN Paris).
Homeomastax jeanninae (Descamps 1971)
Eumastax jeanninae Descamps, Caldasia 11: 139, 158.
Type locality.— Colombia: Dept. Cundinamarca: Caparrapí, 1100 m.
Location of type specimen.— MNHN Paris, holotype male. Female unknown.
Descamps 1979: 136 - transfers to Homeomastax.
Diagnosis and description.— Male subgenital plate as in Fig. 2. The holotype itself was not available to us. From Descamps' (1971) description: “Elytra longer than abdomen, E/F 1.02. Cerci short, conical, straight, very thick at the base, the tip narrowly rounded. Subgenital plate with the upper margin in lateral view curved and angular, the lower margin little convex, except somewhat after the transverse furrow; posterior margin deeply notched in apical view; a very weak posterior carinula. Same type of coloration as in dereixi: vertex and cheeks black, first 2 antennal segments greenish, ventrally brown. The red part of the posterior femur darkened with 2 incomplete rings of brown”.
Male internal genitalia (examined) as in Fig. 14: very similar to those of H. dereixi, to which it is presumably closely related. Epiphallus (Fig. 14C) longer than wide, trapezoidal in dorsal view, narrower at the posterior end, the anterior part containing a large V-shaped membranous region, the posterior margin slightly concave; sensory plaques at base of lophi large. Subepiphallic sac present, short, widening anteriorly. Dorsal fold short and thick, indented medially, mostly hidden under the tightly appressed epiphallus. Sclerite sac large, longitudinally pleated. Ectophallic sclerites more or less straight, the tips slightly diverging, in lateral view broadly triangular with a slightly truncate point (Figs 14A, C). Apical shields simple, forming two more or less vertical plates, lacking a dark pigment spot. Ventroapical evagination and medial apodeme absent. Endophallic plate (Fig. 14C) almost circular at the anterior end, with an unusually large aperture and slot between the arms, only narrowly flanged. Ejaculatory sac (Fig. 14A) large, hanging vertically below end of endophallic plate and not slanted posteriorly. Spermatophore sac simple; its floor curves ventrally at its distal end, well below the level of the arms of the endophallic plate.
Dimensions.— See Table 10.
Distribution.— (Fig. 24) Known only from type locality, Colombia, Dept. Cundinamarca.
Homeomastax brachyptera n. sp.
Holotype.— ♂: PANAMA: Prov. Coclé: Cerro Copé, 1830 m, 20 Sep 1997 (Rowell CHF, Bentos-Pereira A), specimen no. 97398 (ANSP).
Paratypes.— as holotype, 6 ♀ ♀, specimen nos 97392, 97393, 97394, 97395, 97408, 97409; as above, but 15 Sep 1999, 1 ♂, specimen no. 99063 (all ANSP).
Data as above, 1 ♂, specimen no. 97396; three ♀ ♀, specimen nos 97389, 97397, 97492; as above, but 15 Sep 1999, 1 ♀, specimen no. 99052 (MNHNP);
as above, 1 ♂, specimen no. 99057, 1 ♀, specimen no. 99056 (UP).
Other (non-paratypic) material examined.— 1 ♂, data as holotype, specimen no. 97610; 15 Sep 1999, 2 ♂ ♂, specimen nos 99058, 99093, 3 ♀ ♀, specimen nos 99053–99055 (all RC).
Etymology.— Greek brachys, short, and pteron, wing.
Diagnosis.— E/F <0.6, the shortest wings found so far in the genus. Frons predominantly yellow in both living and dried specimens, as opposed to white or blue in other species, and without vermilion marks in female. Posterior margin of pronotum less rounded, more angular, than in other species, forming an angle of approximately 130°.
Male and female subgenital plates as in Figs 2 and 3. Female subgenital plate deeply trilobate, the individual lobes medially grooved and with melanized, ornamented borders. Male subgenital plate similar to that of H. veraguae, but shorter and with a longer medial carina.
Internal male genitalia (Fig. 15). Epiphallus deeply emarginate on posterior margin (Figs 15B, C), tightly appressed to dorsal fold; sensory plaques at base of lophi very long and narrow. Dorsal fold simple, rounded in dorsal view (Figs 15C, G). Subepiphallic sac present, small (Fig. 15D). Ectophallic sclerites long and straight, extending forwards to nearly half the length of the endophallic plate; terminating posteriorly in a straight projection with a rounded tip, narrow in dorsal view and wide and bluntly tapering in lateral view (Figs 15D, G). Sclerite sac present, large, pleated, not clearly delimited ventrally from spermatophore sac. Apical shields simple, decorated laterally with a dense patch of black pigment (Fig. 15D). Endophallic plate (Fig. 15E) rounded anteriorly, almost without lateral flange. Ventroapical evagination absent, but membrane between ventral edges of the apical shields provided with longitudinal folds (Fig. 15F). The phallic complex is very similar to that of the S. American H. bouvieri; it differs from the latter only in its straighter ectophallic sclerites, the very emarginate posterior margin of the epiphallus, and the narrower endophallic plate. H. bouvieri is also larger and fully winged.
Description.— A rather small species with very short wings (Table 11). Sexual dimorphism in size moderate: male/female ratios 0.82–0.87, mean 0.84 (Table 23)Table 23.. Female with 22 outer hind tibial spines and 23 inner ones, male with 22 and 22. Male with white basal antennal segments and frons, the latter yellowish ventral to medial ocellus; clypeus and labrum blue; female with greenish basal antennal segments, face yellow, clypeus and labrum blue.
Distribution.— (Fig. 24) To date known only from the type locality in West-Central Panama, where it is found on both Atlantic and Pacific sides of the watershed.
Natural history.— Clearly associated with ferns, which it was also seen eating. None of the females captured in September had ripe eggs, but copulation was seen.
The kressi species-group
Diagnosis.— Members of this group share the following characteristics:
— Ventroapical evagination large, membranous; if sclerotized then simple, without cuticular teeth or ornament. Medial apodeme present.
— Dorsal fold more or less bifid, with paired sclerotized areas in dorsal surface (except in H. cerciata).
— Endophallic plate large, tending to oblong at anterior end.
— Epiphallic and sclerite sacs present and well developed.
Homeomastax kressi (Rehn & Rehn 1934)
Eumastax kressi Rehn & Rehn 1934: 59.
Type locality.— N. end of Suretka trail, along Duroy River (V. de la Estrella), Prov. Limón, Costa Rica.
Location of type specimen.— ANS Philadelphia.
Descamps 1971: 138, 156.
Descamps 1979, Annls Soc. ent. Fr. (N.S.) 15(1): 136: transfers to Homeomastax.
Diagnosis.— Subgenital plates of male and female as in Figs 2, 3. Female subgenital plate with rather small lateral lobes, a well-marked terminal depression just before the tip, and a few weak marginal teeth; the Panamanian population has better developed teeth. Male subgenital plate long, pointed in both lateral and dorsal view, upper margin in lateral view straight (Costa Rica) or convex (Panama). Male cerci thin, incurved, slightly upwardly inflected at tip in lateral view.
Male internal genitalia (Fig. 16). Epiphallus (Figs 16A–C) long and narrow, tapering anteriorly, with anterior membranous area; sensory plaques small. Subepiphallic sac present. Dorsal fold deeply bifid, with two sclerotized plaques in upper surface (Fig. 16G). Sclerite sac large, pleated. Ectophallic sclerites (Figs 16 C, D, H) robust, spatulate at their tip in lateral view, rounded in dorsal view, with lateral and medial processes at the anterior end. Ventroapical evagination large, membranous, extending dorsally well beyond the ectophallic sclerites; median apodeme present.
Description.— A rather large, long-winged species (Table 12), of moderate sexual dimorphism, male/female ratios 0.80–0.86, mean 0.83 (Table 23)Table 23.. Female with 24 inner and outer hind tibial spines, male with 25 outer and 22 inner spines. Basal antennal segments, face, clypeus and labrum generally whitish; in females the facial carinae are marked with vermilion. Costa Rican specimens are darker, with blue-black pedicel and blackish margins to clypeus and labrum; red portion of femur shadowed proximally and distally with brown. Individuals from the Panamanian end of the species range have somewhat more slender ectophallic sclerites and small differences in the shape of the subgenital plates (Fig. 2); they are also slightly (3–9%) but non-significantly smaller and shorter winged.
Distribution.— (Fig. 25) Lowland Caribbean forest from the Valle de la Estrella (S.E. Costa Rica), via the Ríos Sixaola, Changuinola, Teribé and Guarumo at least as far as Calovébora in West-Central Panama; from sea level to 800 m altitude in valley of the Río Guarumo.
Material examined.— COSTA RICA: Prov. Limón:
3 km SW of Home Creek, 80 m, LS 398800_594100, 2 Jul 1980 (Rowell CHF, Rowell-Rahier M, Hyde C), 1 ♂, 1 ♀, specimen nos 80160, 80460 (RC).
Margarita, trail up Fila, 9.4 km after Bribrí on rd. to Sixaola, 80–205 m, LS 395500_598500, 3 Jul 1980 (Rowell CHF), 1 ♂, specimen no. 80171 (RC).
As above, but 4 Jul 1980, 1 ♂, 3 ♀ ♀, specimen nos 80173, 80544–80546 (RC).
Cahuita, LN 191750_663480, 25 Feb 1988 (Menke S), 3 ♂ ♂, 1 ♀ (USNM).
Cahuita, 0–10 m, LN 191750_663480, 22 Aug 1997 (Rowell CHF Singh I), 1 ♂, specimen no. 97171 (RC).
Cerro Uatsi, above Bribrí, 600 m, LS 396900_579600, 23 Aug 1997 (Rowell CHF Singh I), 1 ♂, 2 ♀ ♀, specimen nos 97179, 97180, 97204 (RC).
Valle de la Estrella: Concepción, 3 km SW, 1 km beyond R. Cerere, 60 m, LN 188000_642000, 18 Aug 1979 (Rowell CHF Rowell-Rahier M), 2 ♂ ♂, 3 ♀ ♀, specimen nos 79004–79007, 79165 (RC).
Valle de la Estrella: Progreso, Finca Cantón, 80–200 m, LN 192300_641000, 27 Jun 1980 (Rowell CHF Rowell-Rahier M Hyde C), 7 ♂ ♂, 3 ♀ ♀, specimen nos 80136, 80147, 80536 to 80543 (RC).
As above but 26 Jun 1980, 1 ♀, specimen no. 80144.
Amubri, 70 m, LS 385000_578100 (Gallardo G) Mar 1995, specimen nos 2-181022, 2-181023, 2-181024; Sep 1996, specimen nos 2-481606, 2-481607, 2-481608; Mar 1996, specimen nos 2-465240, 2-465241, 2-465243; May 1994, specimen nos 1-872129; Apr 95, specimen no 2-144642; 14 Jan 1994, specimen nos 1-844410, 1-844411; 21 Oct 1992, specimen nos 963874, 963876; (Chavarria MM), 14 Jan 1992, specimen no. 2-476119 (all INBio).
Est. Hitoy-Cerere, 100 m, LN 184600_643400 (Carballo G) Sep 1990, specimen no. 0-645554; Jun 1991, specimen nos 434556, 434582; Jun 1991, specimen no. 0-434556; Jul 1991 specimen nos 0-457023, 0-320199; 16 Jan 1992, specimen no. 0-870634; Jul 1992, specimen no. 0-709690; 21 Feb 1993, specimen no. 1-705069 (all INBio).
Other new records.— COSTA RICA: Prov. Limón:
Suretka, 100 m, LS 392150_579500, 20 Sep 1983 (Rowell CHF).
5 km N. of Suretka, trail to Rio Uatsi. 200–220 m, LS 395400_581800, 20 Sep 1983 (Rowell CHF).
Valle de la Estrella: R. Cochey, 230 m, LN 192500_636600, 5 Jul 1980 (Rowell CHF Rowell-Rahier M Hyde C).
PANAMA: Prov. Bocas del Toro.
Isla Colón, 23 Jul 1999 (Cambra R), 5 ♂ ♂, 4 ♀ ♀, 2 ♀ ♀ larvae, specimen nos 99535 to 99545 (UP).
Wekso-Región Teribé, PILA (= Parque Internacional de la Amistad), 20 Oct 1999 (De Gracia L Santos A), 1 ♂, 4 ♀ ♀, 1 ♀ larva, specimen nos 99552 to 99557 (UP).
10 km W. on road from Punta Peña to Almirante, 21.09.99 (Rowell CHF Bentos-Pereira A) 1 ♀, 1 pair in cop., specimen nos 99181–83 (RC)
Rambala, 4 km on trail to Quebrada Platanarito, 150 m, 20 Sep 1999 (Rowell CHF Bentos-Pereira A), 1 ♀, specimen no. 99178 (RC).
Altos del Valle, Río Guabo, 5.5 km after summit on rd. from Fortuna to Chiriquí Grande, 400–500 m, 28 Sep 1997 (Rowell CHF Bentos-Pereira A), 3 ♂ ♂, 4 ♀ ♀ (specimen nos 97575 to 97581 (RC).
Quebrada Felix, 2 km. NW of summit of rd. from Fortuna to Chiriquí Grande, 830 m, Lambert coords. 9723500_365800, 19 Sep 1999 (Rowell CHF, Bentos-Pereira A), two ♀ ♀, specimen nos 99147, 99170; 22 Sep 1999, 1 ♂, specimen no. 99202 (RC).
Prov. Veraguas:
Calovébora; Río Caloveborita, Guazarita, 16 Feb 2000 (De Gracia L Bermudez S), 1 ♂, 1 ♀, specimen nos 2000190, 2000191 (UP).
Homeomastax hondurensis (Rehn & Rehn 1934)
Eumastax hondurensis Rehn & Rehn 1934: 76.
Type locality.— Honduras: Atlantida: Lancetilla, near Tela: Sierra Pija, 700 m.
Location of type specimen.— ANS Philadelphia.
Descamps 1971: 138, 155.
Descamps 1979: 136: transfers to Homeomastax.
Diagnosis.— Subgenital plates of male and female as in Figs 2 and 3. Female subgenital plate long, with well-sclerotized lateral lobes and prominent but sparse marginal teeth, proximally a well-marked medial carina. Male subgenital plate similar to that of H. silvicola with a prominently convex upper margin in side view, but cerci different, short and noticeably thickened at tip.
Male internal genitalia as in Fig. 17. Epiphallus subcircular in dorsal view (Fig. 17E), semimembranous; transparent sensory plaques at base of lophi very large, almost touching in midline. Tips of lophi irregularly truncate in dorsal view, pointed in lateral view (Fig. 17A). Subepiphallic sac well developed. Dorsal fold large and thick, only slightly bifid, but flanked laterally and ventrally by two finger-like evaginations (Figs 17A, D, E). Sclerite sac present (Fig. 17B), longitudinally pleated, extends ventrally to half the length of the ectophallic sclerites. Ectophallic sclerites bifid in lateral view, with almost equally strong upper and lower processes (Figs 17A, E), the lower processes divergent. Ventroapical evagination large, simple, sclerotized towards tip (Fig. 17A); medial apodeme present (Fig. 17E). Endophallic plate (Fig. 17F) of normal dimensions (contra Descamps 1971) and position, anterior end provided with a ridged wing, slightly notched medially.
Description.— A medium sized species with rather short wings (Table 13). Sexual dimorphism in size pronounced: male/female ratios 0.74–0.87, mean 0.80 (Table 23)Table 23.. Male with 26 outer hind tibial spines and 23 inner ones; in the female allotype the only remaining hind femur is broken at the tip. Face discolored in available male specimens; in female, basal antennal segments were probably bluish, face yellow to vermilion, clypeus yellow, labrum blue. No brown bands or shading on the red part of the femur, only the knee itself is brown.
Distribution.— Known only from the type series from the north Caribbean coast of Honduras.
Material examined.— Holotype ♂, allotype ♀ and paratype ♂, data as holotype.
Homeomastax cerciata (Hebard 1923)
Eumastax cerciata Hebard 1923: 178.
Type locality.— Panama: Colón: Porto Bello (=Portobelo).
Location of type specimen.— ANS Philadelphia.
Hebard 1924: 92.
Rehn & Rehn 1934: 79.
Descamps 1971: 138.
Descamps 1979: 136: transfers to Homeomastax.
H. cerciata has many unique characteristics and may well constitute a separate species-group of its own. In the structure of the ventroapical evagination it is most similar to the kressi species-group, and is therefore included tentatively here.
Diagnosis.— Subgenital plates of male and female as in Figs 2, 3. Female subgenital plate rather short and blunt with a short obtuse point, most of surface darkly pigmented, margin devoid of teeth. Male subgenital plate similar to that of H. robertsi, with a convex but sinuate upper margin in side view, but more carinate medially. Male cerci unmistakable, large, heavily built, abruptly deflected downwards by 60° at half-length.
Male internal genitalia (Fig. 18). Epiphallus with slightly concave posterior margin, body of epiphallus with large V-shaped membranous area, giving at first the impression of a deeply incised anterior margin (Fig. 18E). Subepiphallic sac present, wide and shallow, providing an unusually mobile link between phallus and epiphallus. Dorsal fold unitary, oblong, very short, without sclerites, the posterior border provided with numerous fine denticles (Figs 18A, D). Sclerite sac present, simple, long, extending almost to anterior margin of spermatophore sac and joined to it in the midline, but the two cavities are not confluent (Figs 18B, C). Ectophallic sclerites in lateral view straight, truncate at anterior end, ending posteriorly in a sharp downwardly inflected point (Figs 18A, B); in dorsal and apical view remarkable, with parallel rows of fine teeth following the contours of the subapical regions (Fig. 18G); in dorsal view the posterior ends of the sclerites first converge, then diverge at the tips (Fig. 18D), which are spatulate in dorsal view. Spermatophore sac opens in central portion of genital trough; posteriorly the sac extends two free finger-like processes on either side of the genital trough, visible beneath the posterior ends of the ectophallic sclerites (Figs 18C, G), a unique feature. Ventroapical evagination large, membranous, projecting posteriorly rather than dorsally, inserting dorsally on the underside of the endophallic plate some distance from its tip (Figs 18B, C); medial apodeme present, but not bifid as usual. A low ridge of membrane runs internally along the medial surface of the ventral ectophallic membrane from the medial apodeme to a small sclerite at the anterior edge (Fig. 18A); the entire ventral surface of the ectophallus is lightly sclerotized to this point. Endophallic plate rounded-oblong in outline anteriorly, with a ribbed flange surround (Fig. 18H).
Description.— One of largest species of the genus, long winged (Table 14). Sexual dimorphism relatively small, male/female ratios 0.87–0.91, mean 0.89. Hind tibial spines (female), 23 outer and 24 inner, 23 and 21 in a male. Basal antennal segments, frons, genae, clypeus and labrum whitish in both sexes when dried. Red portion of femur shadowed proximally and distally with brown.
Distribution.— (Fig. 25) To date known only from Portobelo (the type series) and from Cerro Campana, both in Central Panama. The species found in the Atlantic lowlands near Portobelo is however H. veraguae; the type series of H. cerciata therefore probably came from higher ground in the modern Chagres National Park behind Portobelo, or conceivably was mislabelled. The collector (A. Busck) also collected near Cerro Campana.
Material examined.— PANAMA: Prov. Panamá: Cerro Campana, 900 m, trail through forest, 7 Sep 1997 (Rowell CHF & Windsor D), 5 ♂ ♂, 1 ♀, specimen nos 97298 to 97302, 97600 (RC). Same data, but 13 Sep 1997 (Rowell CHF), 6 ♂ ♂, 1 ♀, specimen nos 97601–97607; same data, but (Sanchez A), 2 larvae, specimen nos 97608, 97609 (all UP).
Natural History.— On ferns in montane forest.
Homeomastax tenoriensis n. sp.
COSTA RICA: Guanacaste: Volcan Tenorio: near summit of road from Tierras Morenas to Bajo Los Cartagos, LN 287250_426500, 1040 m, 21.7.1991 (Rowell CHF Elsner N), holotype ♂, specimen no. 91103 (ANSP).
Paratypes: one ♂, two ♀ ♀, data as above, specimen nos 91104–91106 (ANSP).
Guanacaste: Tilarán, Z.P. Tenorio A.C.A., Río S. Lorenzo, 1050 m. LN 287800_427600, May 1994 (Rodriguez G), one ♀, specimen no. CRI001 873903 (INBio).
Etymology.— specific name derived from the type locality.
Diagnosis.— Subgenital plates of male and female as in Figs 2, 3. Female subgenital plate intermediate between those of kressi and cerciata, lateral lobes weakly sclerotized and not prominent, margin devoid of teeth. Male subgenital plate short, with a faint medial carina but no medial notch; in lateral view the posterior surface forms a right angle with the straight dorsal rim; in dorsal view smoothly rounded with a minute apical lip. Male cercus slightly inwardly incurved, with bluntly rounded, slightly swollen tips.
Male phallic complex (Fig. 19). Epiphallus oblong, parallel sided, with a straight posterior margin between moderately large sensory plaques; anterior margin with a U-shaped membranous area (Fig. 19B). Subepiphallic sac present. Dorsal fold (Fig. 19D) only slightly bifid, with two sclerotized plaques in upper surface. Ectophallic sclerites (Figs 19D, F) simple, spatulate at their tip in lateral view, without process at the anterior end; apical shields small. Sclerite sac present, longitudinally pleated (Fig. 19F). Ventroapical evagination (Figs 19D–G) large, membranous, clearly divided into two parts: an upper lobe which forms an inverted hood over the end of the genital trough, provided with fine chitinous teeth, and a lower spoon-shaped lip; median apodeme present.
Description.— Somewhat smaller than Costa Rican kressi; markedly more brachypterous (Table 15) and slightly more sexually dimorphic in size, male/female ratios 0.79–0.88, mean 0.82 (Table 23)Table 23.. Outer hind tibial spines of female 21, inner tibial spines 20; in male, 19–20 for both. Male face white, basal segments of antennae and clypeus and labrum, blue; in female all greyish white, face marked with vermilion. Red portion of femur with heavy brown shading proximally and distally. General coloration rather dull for the genus.
Distribution.— (Fig. 25) Known to date only from montane forest on the southern slopes of Volcan Tenorio, the southernmost peak of the Cordillera de Guanacaste.
The silvicola species-group
Diagnosis.— Members of this group share the following characteristics:
— Ventroapical evagination large and complex, appearing chelate in lateral view, lightly sclerotized and provided on the folded surface with small chitinous teeth; medial apodeme large.
— Ectophallic sclerites with a barbed dorsoapical crest, and ending apically in palmate claw-like structures.
— Dorsal fold simple or bilobed, provided with paired sclerites.
A very distinctive and homogenous group of closely related species from northern Costa Rica.
Homeomastax silvicola (Rehn & Rehn 1934)
Eumastax silvicola Rehn & Rehn 1934: 72.
Type locality.— La Emilia, nr. Guápiles, Costa Rica, 1000′.
Location of type specimen.— ANS Philadelphia.
Rehn 1905a: 675 as Eumastax dentata (misidentification).
Descamps 1973b: 138.
Descamps 1979, Annls Soc. ent. Fr. (N.S.) 15(1): 136: transfers to Homeomastax.
Diagnosis.— Subgenital plates of male and female as in Figs 2 and 3. Female subgenital plate shorter and more slender than in robertsi, margin devoid of teeth, lateral lobes less developed. In both species there is a well-marked transverse fold or depression just anterior to the shallow lateral lobes. Male subgenital plate in lateral view with a smoothly convex dorsal margin (not sinuate as in robertsi), similar to that of H. hondurensis; cerci short and smoothly tapered.
Male phallic complex (Fig. 20). Epiphallus more or less square in dorsal view, sensory plaques rather small, posterior margin straight, lophi long and thin. Subepiphallic and sclerite sacs present, but poorly developed. Dorsal fold (Figs 20E, J) short, convex, bilobed, with pointed lobes; two sclerotized plaques in upper surface. Internal surfaces of ectophallic sclerites straight; sclerite with a large dorsal barb, oval in dorsal view; terminates in three sharp ventrally directed points, like a clawed foot (Figs 20E, J). Apical shields rather narrow. Spermatophore sac opens only to the posterior region of the genital trough. Ejaculatory sac small. Ventroapical evagination as in species-group diagnosis; medial apodeme present.
Description.— A medium sized species with no especially extreme measurements or proportions (Table 16). Sexual dimorphism moderate, male/female ratios 0.82–0.89, mean 0.85 (Table 23)Table 23.. Female with 24 inner and outer hind tibial spines, male with 23 and 23. Frons whitish in dried specimens, in female marked with vermilion on carinae, in male bluish ventrally; basal antennal segments, clypeus and labrum, blue.
Distribution.— (Fig. 26) Northern foothills of the Cordillera Central and of the N.E. Talamancas, Costa Rica, from 300–900 m. Not to date recorded from the lowland plain.
Material examined.— COSTA RICA:
Prov. Alajuela: nr Virgen del Socorro, Sarapiquí, 1000–800 m, LN 248800_517500, 22 Jun 1980 (Rowell CHF Rowell-Rahier M Hyde C), 3 ♂ ♂, 2 ♀ ♀, specimen nos 80037–80041 (RC).
Prov. Alajuela: Laguna Hule, 780–730 m, LN 253900_513700, 16 Jul 1991 (Rowell CHF), 1 ♂, specimen no. 91074 (RC).
Prov. Cartago: Bajo Pacuare, 580–700 m, LN 207400_589500, 7 Jul 1980 (Rowell CHF Rowell-Rahier M Hyde C), 1 ♀, specimen no. 80684 (RC).
Prov. Cartago: Juan Viñas, LN 208700_564300, Mar 1902 (L. Bruner), 1 larva, 1 ♂ (ANSP).
Prov. Cartago: Turrialba, LN 21000_571000, 22 Feb–18 Mar 1965 (Duckworth SS & WD), 5 ♂ ♂, 1 ♀ (USNM).
As above, but 2 Jun 1973 (Ekis G), 1 ♂ (USNM).
As above, but 25 Mar 1953 (Cartwright OL), 3 ♀ ♀, 2 ♂ ♂ (USNM).
Prov. Cartago: Monumento National Guayabo, 1100 m, LN 217400_570000, 15 Jul 1992 (Fonseca G), specimen no. CRI001 888239 (INBio).
Prov. Cartago: Grano De Oro, 1120 m, LN 200250_595900 14 Apr 1993 (Campos P) specimen nos CRI001 307246 & CRI001 307248 (INBio).
Prov. Limón: Río Toro Amarillo, crossing with rd. from Guápiles, 305 m, LN 243300_556400, 9 Jun 1980 (Rowell CHF Rowell-Rahier M Hyde C), 4 ♂ ♂, 3 ♀ ♀, specimen nos 80042–80048 (RC).
Prov. Limón: 3 km N. of Carrillo: Finca El Tapir, 550 m, LN 236000_542000, 13 Jul 1991 (Rowell CHF Elsner N), 1 ♂, specimen no. 91074 (RC).
Prov. Limón: Río Toro Amarillo, 7 km N of Guápiles, 0.5 km N of Quebrada Grande, 550 m, LN 237300_558100, 14 Jul 1991 (Rowell CHF & Elsner N), 2 ♂ ♂, specimen nos 91047, 91048 (RC).
Prov. Limón: Río Toro Amarillo, 10 km N of Guápiles, S of Quebrada Grande on trail to S. Valentino. 650 m, LN 234000_558400, 10 Sep 1993 (Rowell CHF), 1 ♂, 1 ♀, specimen nos 93209, 93210 (RC).
Prov. S. José: Bajo La Hondura, 900 m, LN 227300_538300, 31 Jul 1991 (Rowell CHF & Elsner N), 1 ♀, specimen no. 91206 (RC).
Other new records.— Prov. Cartago: nr Chitaría, 590 m, LN 214000_582000, 24 Jun 1980 (Rowell CHF Rowell-Rahier M Hyde C).
Prov. Cartago: Turrialba, 3 km SE, grounds of CATIE, forested river gorge. 500 m, LN 208500_575200, 17 Feb 1973 (Rowell CHF).
Other older records (ANSP) included on distribution map.— Carrillo; Guácimo; Peralta; Chitaría; Pejivalle; Navarro; Guápiles.
Homeomastax robertsi (Descamps 19739)
Eumastax robertsi Descamps 1973b: 953.
Type locality.— Costa Rica: Prov. Heredia; Sarapiquí, Puerto Viejo, Finca La Selva.
Location of type specimen.— ANS Philadelphia.
Descamps 1979: 136: transfers to Homeomastax.
Diagnosis.— Subgenital plates of male and female as in Figs 2 and 3. Female subgenital plate somewhat longer than in H. silvicola and with wider lateral lobes, margin sparsely toothed. Descamps' (1973b) figure is of an extreme example — the subgenital plate is shorter in most individuals. Male subgenital plate with upper margin in lateral view convex but sinuate, the apex more acute than in H. silvicola; this is the best character to distinguish this species.
Male phallic complex (Fig. 21) very similar to that of silvicola — crest and barb on the ectophallic sclerites (Figs 21 A, B) slightly higher and narrower, and the claw-like ending of the sclerite somewhat less pronounced. Dorsal fold (Fig. 21B) longer and narrower, bilobed. Lophi of epiphallus truncate in axial view (Fig. 21C), sharp in lateral view (Fig. 21A).
Description.— Slightly larger than E. silvicola in the female, and with fractionally longer wings (Table 17). Sexual dimorphism moderate, male/female ratios 0.79–0.87, mean 0.84 (Table 23)Table 23.. Hind tibial spines (female), 26 outer and 21 inner, male with 22 and 23. Male with basal antennal segments and frons blue, clypeus and labrum dark blue; female with blue pedicel, scape vermilion proximally, blue distally, frons vermilion, clypeus and labrum blue.
Distribution.— (Fig. 26) Northeastern Costa Rica and southeastern Nicaragua, from sea level to 1000 m. There is no clear boundary separating the upper part of its zone of occurrence from that of H. silvicola. The two female paratypes from Turrialba (Descamps 1973b: 955 - not examined) are probably misidentified H. silvicola, as all other specimens from that locality belong to that species.
Material examined.— COSTA RICA:
Prov. Alajuela: S. Lorenzo, some km S, 700 m, LN 221550_519600, 5 Sep 1976 (Rowell CHF), 1 ♀, specimen no. 76057 (RC).
Prov. Alajuela: Dos Ríos: Finca Aurora 700–800 m, LN 320000_385000, 12 Oct 1999 (Rowell CHF), 2 ♂ ♂, 1 ♀, specimen nos 99463–99465 (RC).
Prov. Alajuela: Dos Rios: Est. S. Ramón Oeste, 620 m, LN 318100_381900, 14.04.1994 (Moraga C), one ♂, specimen no. CRI001 779905 (INBio).
Prov. Heredia: Puerto Viejo: Finca La Selva, 40 m, LN 268800_535300, 27 Mar 1986 (Rowell CHF), 2 ♂ ♂, 1 ♀, specimen nos 86454, 86456, 86031(RC).
As above, 29 Mar 1986, 1 ♀, specimen no. 86457 (RC).
As above, 30 Jan 1973, copulating pair, specimen nos 73014, 73015 (RC).
As above, 30 Jan 1978, 1 ♀, specimen no. 78044 (RC).
As above, 11 Sep 1979, (Rowell CHF, Rowell-Rahier M), 1 ♀ specimen no. 79234 (RC).
As above, 15 Jul 1991, (Rowell CHF Elsner N), 1 ♀, specimen no. 91080 (RC).
Prov. Heredia: Cantarrana-Quebrada Negra, LN 261000_531400, 1 Apr 1986 (Rowell CHF), 1 ♀, 1 ♂, specimen nos 86470, 86032 (RC).
Prov. Heredia: Ridge between R. Peje & R. Sardinalito, 700 m, LN 253600_527800, 7 Apr 1986 (Rowell CHF), 1 ♂, specimen no. 86483 (RC).
Prov. Heredia: Sarapiquí, ridge between R. Peje & R. Sardinalito, 960 m, LN 250650_527900, 9 Apr 1986 (Rowell CHF) (RC).
Prov. Heredia: Sarapiquí: Finca Tirimbina, LN 264650_524550, 23 Feb 1973 (Rowell CHF), 1 ♀, specimen no. 73013 (RC).
Prov. Heredia: Sarapiquí: La Virgen, 220 m, LN 264600_521500, 16 Apr 1993 (Ortiz M), specimen no. CRI001 300731 (INBio).
Prov. Heredia: Sarapiquí: Est. Magsasay, 200 m, LN 264600_531100, 15 Jun 1990 (Alcazar E), specimen no. CRI000 222646 (INBio).
Prov. Limón: PN Tortuguero: Est. Cuatro Esquinas, 10 m, LN 280000_590500, 16 Apr 1989 (Aguilar R), specimen no. CRI000 83020 (INBio).
Prov. Limón: Cerro Tortuguero, 120 m, LN 285000_588000, 16 Mar 1993 (Delgado R), specimen no. CRI001 352892 (INBio).
Prov. Limón: PN Tortuguero: Sector Cocori II, 150 m, LN 286000_567500, 20 Sep 1992 (Rojas E), specimen no. CRI000 976283 (INBio).
Prov. Limón: Río Sardinas, Barra del Colorado, 50 m, LN 291500_564700, 06 Sep 1994 (Araya FV), specimen no. CRI002 030592 (INBio).
NICARAGUA: Zelaya, El Recreo (forêt). 30 m, 12° 10′N, 84° 19′ E, 15 Oct 1984 (Amédégnato C, Poulain S), 1 ♂, 1 ♀ (MNHNP).
Homeomastax strigla n. sp.
COSTA RICA: Prov. Guanacaste: P.N. Rincón de la Vieja, fila above R. Colorado, 930 m, LN 307000_389500, in tree-fall clearing in forest, 20 Jul 1980 (Rowell CHF & Rowell-Rahier M), holotype ♂, specimen no. 80547 (ANSP).
Etymology.— Latin strigla, witch, noun in apposition, an allusion to the Spanish name of the type locality, translatable as the witch's nook, probably because of the steaming fumaroles found there.
Diagnosis.— Subgenital plates of male and female as in Figs 2 and 3. Female subgenital plate similar to that of silvicola, but yet more slender and with longer, narrower, ornamented margins. Male subgenital plate very distinctive — short in lateral view, with a large vertical lip medially; medially carinate to a variable extent.
Male internal genitalia (Fig. 22): epiphallus with emarginate anterior border, lophi thin and very sharp; posterior margin concave, sensory plaques oblong in dorsal view, rather than domed, small (Fig. 22D). Subepiphallic sac present, broad, shallow. Dorsal lobe unitary, with two large thin sclerotized areas that do not extend to the posterior edge (Fig. 21A). Sclerite sac large, with lateral projections, complexly folded. Ectophallic sclerites (Figs 22 A, B, G) of same type as in silvicola and robertsi, but more slender, with a higher crest and a narrow barb, terminating in one blunt process and a single lateral claw (i.e., it terminates in two processes, not three, as in the other species); at anterior end sclerites outward curving. Ventroapical evagination large, similar to those of robertsi and silvicola. Endophallic plate oblong and winged at anterior end, upwardly curved at posterior end. Spermatophore sac opening in posterior half of genital trough.
Description.— One of the largest species of the genus, only H. cerciata is comparable (Table 18). Sexual dimorphism moderate, male/female ratios 0.80–0.91, mean 0.86. Outer hind tibial spines, 24 (male), 26 (female), inner spines, 21. Frons white in dried specimens, in female tinged vermilion on carinae; basal antennal segments, clypeus and labrum blue in male, greyish white tinged blue in female.
Distribution.— (Fig. 26) Confined to montane forest of the northern peaks of the volcanic Cordillera de Guanacaste and the southern part of the Cordillera de Tilarán, between 500 and 1400 m. Replaced on V. Tenorio by H. tenoriensis n. sp. (see above). As discussed under H. dentata below, that species and H. strigla may in fact be synonymous, in which case the range of the species would be extended somewhat to the south.
Material examined.— Holotype ♂, as above.
Paratypes, as holotype, but 2 ♂ ♂, specimen nos 80259, 80548, 1 ♀, specimen no. 80549 (all ANSP).
COSTA RICA: Prov. Alajuela: La Libertad, in valley of R. Caño Negro, between P.N. Rincón de la Vieja and Piedras Blancas, 400–500 m, LN 312000_413000, 21 Jul 1980 (Rowell CHF & Rowell-Rahier M), 1 ♀, specimen no. 80685 (ANSP).
Prov. Guanacaste: S.E. slope of Volcan Cacao, 1040 m, LN 323300_375300, 24 Jul 1991 (Rowell CHF Elsner N Chavez C), 1 ♂, 1 ♀, specimen nos 91155, 91156 (MNHNP).
Prov. Guanacaste: R. Naranjo, 3 km SE: Hca Montezuma, 500 m, LN 293000_418000 (Parker FD), 1 ♂, specimen no. 94072 (SUUL).
Non-paratypic material.— Prov. Guanacaste: Volcan Miravalles: Sitio La Gigante, 1080 m, LN 297100_413150, 20 Jul 1991 (Rowell CHF & Elsner N), 1 ♂, specimen no. 91090 (RC).
Prov. Guanacaste: S.E. slope of Volcan Cacao, 1040 m, LN 323300_375300, 24 Jul 1991 (Rowell CHF Elsner N Chavez C), 2 ♂ ♂, spec. nos 91151, 91152, 2 ♀ ♀, specimen nos 91153, 91154 (RC).
As above, but Sep 1991 (Chaves C), 1 ♂, spec. no. CRI000 331708 (INBio).
Prov Guanacaste: P.N. Guanacaste: Estac. Pitilla, 7.5 km S of Sta. Cecilia, 700 m, LN 330200_380400, Feb 1990 (Rios P Moraga C Blanco R), 2 males, specimen nos CRI000 175945, CRI000 175889 (INBio).
As above, but Apr 1992 (Rojas S), 1 ♀, specimen no. CRI000 418279 (INBio).
Prov. Alajuela: Colonia Palmareña, 700 m., 9 km SO de Bajo Rodriguez, LN 245900_475900. March 1997 (Carballo G), 1 pair in cop., specimen nos CRI002 537403 & 537404 (INB io).
Prov. Alajuela: Reserva biológica S. Ramón, 800 m, LN 244100_472100, 28 Oct 1994 (Carballo G), one ♂, specimen no. CRI002 109670 (INBio).
As holotype, but 80548 (in alcohol, ♀ dissection) (RC).
Species of as yet unknown affiliation
Homeomastax dentata (Saussure 1903)
Eumastax dentatus Saussure 1903: 91.
Type locality.— Costa Rica: Palmares, near San Ramón, at head of R. Grande de Tárcoles, c. 3000 feet.
Location of type specimen.— holotype ♀: MNHN Paris.
Kirby 1910: 77 as Eumastax dentata (correction of gender).
Rehn & Rehn 1934: 70 (redescription and allocation of male, in error — probable misidentification of H. bustum n. sp.).
Descamps 1971: 139 (in error, following Rehn & Rehn 1934).
Descamps 1973b: 949 (in error, misidentification of H. bustum n. sp.).
Descamps 1979: 136 (transfers to Homeomastax).
Rowell et al. 1983: 207 (ecology — in error, following Rehn & Rehn (1934) and Descamps (1971), misidentification of H. acrita n. sp.).
Rowell 1998: 226 (distribution — in error, misidentification of H. acrita n. sp.)
See remarks under H. surda concerning the description of the putative males of surda and dentata by Rehn & Rehn (1934) and their failure to examine the original types. Examination of the holotype of H. dentata shows that this is not the same species as the “dentata” of Rehn & Rehn and all later authors. In size and in morphometric ratios (Table 19) it is many standard deviations removed from those forms. The morphometric ratios of the type fit exactly those of H. silvicola Rehn & Rehn 1934, and slightly less well those of H. strigla n. sp. and of H. bustum n. sp., but correspond to no other species (see Table 23Table 23.). The female subgenital plate (Fig. 3) is compatible with all these identifications. In size, however, the holotype of dentata is larger than any known specimen of any of these three species, though H. strigla approaches it. The type locality of dentata is close (about 30 km) to known modern localities of strigla, and further from those of the other species (Figs 23, 26). Unfortunately the area is now urbanised, and new topotypic examples are not likely to be found. In the absence of topotypic material which includes females of the same size as the type, dentata cannot be redescribed. If a later described species is in fact synonomous with it, this is most likely to be H. strigla n. sp.
Comment.— The male which Rehn & Rehn (1934) assumed to be H. dentata (COSTA RICA: Prov. San José: S. slope of Cerros de Bustamente, between R. Colorado and the Sabanillas de Pirrís, 1500–3940 feet, LN 192900_526900, 25 Aug 1927 (Rehn JAG) (ANSP) is no longer present in the Philadelphia collection, and apparently lost; it cannot therefore be stated with certainty which species they described. Their drawing of its subgenital plate indicates H. bustum n. sp. or H. acrita n. sp., which can be separated with certainty only by the internal male genitalia; the former species is geographically more probable (see Fig. 23). Their putative female (COSTA RICA: Prov. San José: Cangrejal de Aserrí, 800 m, LN 194400_512500, Apr 1906 (Biolley P) (ANSP)) has been examined; it appears to be H. bustum (shape of subgenital plate, dimensions and morphometrics; the elytron is slightly shorter than in the type series) and is certainly not H. acrita. The area around Cangrejal is now largely deforested and probably much drier than in Rehn's time; we have so far been unable to find a modern Homeomastax there.
Distribution.— Known only from type locality in West-Central Costa Rica (Fig. 26).
Homeomastax richteri Descamps 1971
Eumastax richteri Descamps, Caldesia 11: 154.
Type locality.— Colombia: Dept. Sur Santander: Alto Río Opón, 900 m.
Location of type specimen.— MNHN Paris, holotype ♀. Not examined. ♂ unknown.
Descamps 1979: 136 (transfers to Homeomastax).
Description.— (translated from Descamps 1971). “A large species of the surda group” - i.e., Homeomastax as currently understood. “Supraclypeal triangle relatively well-marked, less than 2 times as wide at its base as high. Posterior part of the pronotal disc clearly elevated in lateral view. Elytra projecting beyond the ovipositor, E/F 1.07. Subgenital plate very characteristic, strongly projecting and narrowed towards the rear, trilobed and armed with irregular long spines; medial lobe narrow and much longer than the lateral lobes. Valves of the ovipositor like all the other species. Two pale rings around the base of the posterior tibia.
“Cephalic capsule discoloured. Pronotum greenish yellow; a small dark mark at the superior posterior angle of the lateral lobes; thoracic pleura like the pronotum. Subgenital plate marked with brown. The apical half of the femur bright red, without brown marks; external face of the basal half with brown chevrons (possibly postmortem effects?). Base of hind tibia with 2 incomplete pale rings”.
Dimensions.— see Table 20.
Distribution.— (Fig. 24) Colombia, Sur Santander (known only from type locality).
The close similarity of the female subgenital plate (Fig. 3) to those of bouvieri and brachyptera, strongly suggests the bouvieri species-group, though in absence of a male this cannot be concluded with certainty.
Homeomastax annulipes Descamps 1973
Eumastax annulipes Descamps 1973b: 946.
Type locality.— Colombia: Dept. El Valle, Anchicaya, 300 m.
Location of type specimen.— MNHN Paris, holotype female. Not examined. Male unknown.
Descamps 1979: 136 (transfers to Homeomastax).
Description.— (translated from Descamps 1971) “A medium size species, of the surda group” - i.e. Homeomastax as currently understood. “Posterior margin of pronotal disc excurved. Ends of elytra attain end of abdomen: E/F = 0.89. Elytra little widened posteriorly.
“Subgenital plate very long, a little tapered towards the tip, slightly narrowed a little after the posterior edge of the 8th tergite, the extremity very broadly rounded, covering the basal parts of the ovipositor valves. Posterior edge armed with numerous small spines, pointing dorsally and invisible ventrally; a small triangular lobe in the midline.”
“Face and cheeks pale; vertex with a large black spot; a small black spot behind the eye. Lateral lobes of pronotum with a black band in upper part. Thoracic pleura yellow. Abdomen discolored. Hind femora yellow-green basally, distal half with a large red ring between 2 black rings. Hind tibiae dark brown towards the tip; a pale pregenicular ring.”
“The ringed femora are unique for the group”.
The female subgenital plate is not trilobed and so not obviously reminiscent of the bouvieri species-group, which accounts for most of the other known S. American species of the genus. However, H. dereixi also has a simple subgenital plate, so membership of this group is not excluded.
Dimensions.— see Table 21.
Distribution.— (Fig. 24) Colombia, El Valle (known only from type locality).
Homeomastax sp. indet.
Material examined.— COLOMBIA: Chocó: El Tambo, 5 Apr 1918 (M.A. Carriker), 1 ♀; Boca Murindo, 9 Apr 1918 (M.A. Carriker), 1♀ (both ANSP).
In the absence of a male, the species cannot usefully be described or named. The female subgenital plate (Fig. 3) is unlike that of any known S. American species, resembling most closely those of the silvicola species-group from Costa Rica.
A very large species, F (female) = 15.32 mm, with long wings, E/F = 1.0 (Table 22). Basal segments of antennae, frons, clypeus and labrum all yellowish white. Red portion of femur slightly shadowed with brown proximally, distally clear.
Distribution.— (Fig. 24) Colombia, Chocó.
Discussion
Although Homeomastax now includes 22 species, the genus is probably considerably larger than this. In Costa Rica, which is the best collected Central American country, Homeomastax — like several acridid genera, e.g. Rhachicreagra (Jago & Rowell 1981, Rowell & Amédégnato 2000), Drymophilacris (Rowell 2000) — shows a pattern of a few widely distributed, lowland species, plus numerous local species in the mountains. Given how poorly collected are the other Central American countries and Colombia and Ecuador, it is highly probable that there remain numerous species of Homeomastax to discover. In view of the extensive modern deforestation of much of this area, it is also quite probable that the genus has recently contained “centinelan” local species, made extinct before their description. There seems no obvious reason why the genus should have its true northern limit in Honduras, as suitable environment continues at least to Belize and Guatemala, possibly Mexico. There are no obvious differences in ecology between the species, and the different species are virtually never sympatric, further suggesting that they are not ecologically differentiated. Presumably speciation is largely allopatric and due to distance or spatial boundaries.
Homeomastax is a “difficult” genus — the numerous species are distinct enough in e.g., their internal male genitalia, that there can be little doubt as to their validity, but at the same time they are unusually difficult to distinguish on external characters, at least with the naked eye. A few species, e.g., H. bustum n. sp. & H. acrita n. sp., are indeed only identifiable with certainty on the basis of the internal male genitalia. This raises the question as to why the external morphology is so uniform and conserved and yet male genital morphology apparently very plastic; we have no answer to offer.
Related to this is the question of the age of the species. The Eumastacidae s.l. (i.e., all Eumastacoidea other than the Proscopiidae) are extremely old (>200 million years) and in general show a highly conserved morphology, but southern Central America is very young — the Isthmus is only about 3 million years old. Are we dealing with a recent species radiation, or with ancient South American species which have emigrated into new regions? Molecular systematics, which might answer this question, has to date provided only the information that the split between the Temnomastacinae (an exclusively South American subfamily of the Eumastacidae s.s.) and the Eumastacinae is relatively ancient, probably occurring after only about one third of the evolution of the Eumastacidae s.l., and the divergence between Eumastax and Homeomastax is more recent, but also of considerable age, certainly not a recent event (Matt 1998).
Biogeographically, the silvicola species-group appears predominantly northern, the bouvieri species-group southern, the kressi species-group widely distributed on the Caribbean slope, and the surda species-group widely distributed on the Pacific slope. Exceptions within the surda group are H. veraguae, which extends right across the Panamanian Isthmus to the Caribbean coast, and H. pluvialis, which is a few kilometres onto the Caribbean side of the watershed. The bouvieri species-group extends north in the shape of H. brachyptera to Central Panama. To date we have few specimens of the genus from Eastern Panama, either Darién or the Comarca Kuna; it will be interesting to see whether there are intermediate bouvieri-group species there.
Acknowledgments
We thank Dr. C. Amédégnato (Museum national d'histoire naturelle, Paris), Dr. U. Aspöck and Prof. A.P. Kaltenbach (Naturhistorisches Museum, Vienna), D. Azuma (Academy of Natural Sciences, Philadelphia), Dr. W.J. Hanson (State University of Utah at Logan), Dr. D. Nickle (U.S. National Museum, Washington), Dr. P.D. Perkins (Harvard University Museum of Comparative Zoology), Prof. D. Quintero (Universidad de Panamá), A. Solís (INBio, Costa Rica) and Drs. P. Schwendinger & S. Hauser (Museum d'Histoire Naturelle, Geneva) for access to their collections and/or the loan of specimens. We are also grateful to Drs. C. Amédégnato and Prof. C.S. Carbonell for discussion, Prof. D. Quintero for assistance with Panamanian place names and logistics, numerous colleagues over the years for assistance in field collection, the authorities of Costa Rica and Panama and many private landowners there for permission to collect in National Parks and elsewhere. An anonymous reviewer made very useful comments. A. Bentos-Pereira thanks the Comisión Sectorial de Investigación Científica de la Universidad de la República, Uruguay, for financial assistance.
Literature Cited
Fig. 2.
Homeomastax spp. Male subgenital plates in lateral and axial view. Based on Descamps 1971, with modifications and additions. All drawings except that of H. jeanninae have been compared with specimens. Scale bar, 1 mm.
Fig. 3.
Homeomastax spp. Apices of female subgenital plates in ventral view. The figures of bouvieri, dereixi, richteri and annulipes are taken from Descamps 1971, 1973b; for the first three of these species no scale was provided. All drawings except those of H. richteri and annulipes have been compared with specimens. Scale bar, 1 mm.
Fig. 4.
Homeomastax surda (Burr). General anatomy of the male internal genitalia. In this, as in all other figures, posterior is to the right (in lateral views) or downwards (in dorsal or ventral views). Scale bars, 1 mm. A. Intact phallic complex as normally dissected and figured, without musculature. The dorsal and ventral folds, sensu Descamps 1971, are shown. B. As A, but with ectophallic membrane positioned as in life. The cut edge of the membrane attaches to the rim of the subgenital plate. The “ventral fold” is now seen to be the point at which the membrane reverses and runs posteriorly. All structures anterior to the ectophallic membrane and lateral to its folded portion, including the epiphallus anterior to the lophi, are bathed in haemolymph. C. Parasagittal section at level of the ectophallic sclerite. The preparation has been extended dorsoventrally in order to separate the sacs. The ectophallic membrane is invaginated in the midline to give a subepiphallic sac below the epiphallus, evaginated to give the dorsal fold, and invaginated beneath the dorsal fold to form the sclerite sac, which is continuous with the genital trough. The epiphallus itself is apparently also an invagination of the membrane, more flattened and more sclerotized than the others. D. As B, but in dorsal view. E. Axial view of phallic complex from the rear. The ectophallic membrane has been displaced ventrally to allow a clear view. In H. surda the apical evagination is very small and has no apodeme attached to its base. F. Posterior tip of phallic complex from beneath, with most of ectophallic membrane removed, to show the insertion of the “legs” of the endophallic plate inside the apical shields. G. Endophallic plate, to show the sclerotized anterior portion, and the lightly sclerotized or unsclerotized extensions to the “legs” which fuse with the inner surface of the apical shields. H–K. Epiphalli, dorsal view, from four different preparations, to show variation. Geographical sources: H, Carara; I, Agua Buena; J & K, Pozo Azul de Pirrís.
Fig. 5.
Copulation in Homeomastax. (Scale bars all 1 mm: A, bar above; B,C, bar below; D,E, bar above; F, not to scale. A. Homeomastax strigla n. sp. Copulating pair from right hand side, diagrammatic. Male shaded. B. As A, but showing detail of abdominal contact from left hand side. C. As B, but after relaxing and removing the male, to show extruded and rotated female reproductive aperture. SAP, supra-anal plate; VOV, ventral ovipositor valves; ApB, aperture of bursa copulatrix; SGP, subgenital plate. D. H. robertsi Descamps (see also Fig. 21). Phallic complex with emplaced spermatophore immediately before copulation. It is thought that the pointed end of the spermatophore (arrow) is placed in the aperture of the bursa copulatrix shown in C, when the animals are in the position shown in A & B. E. H. strigla n. sp. Phallic complex dissected from the copulating pair shown in A–C. Note extended position of the ventroapical evagination (arrow); compare with normal position shown in Fig. 22B. F. As E, perspective sketch from above, posterior and to the right (not to scale). Compare with Fig. 21H.
Fig. 6.
Homeomastax carrikeri (Hebard), male genitalia. A. Entire phallic complex, lateral view. B. Parasagittal section through the central region of A at the level of the ectophallic sclerite, to show relation of endophallic plate and apical shield. C. Sagittal section through the central region of A, to show ejaculatory and spermatophore sacs, ventroapical process and its medial apodeme. D. Tip of phallic complex, dorsal view. E. Epiphallus, dorsal view. F. Epiphallus, axial view. G. Endophallic plate, ventral view.
Fig. 7.
Homeomastax veraguae (Hebard), male genitalia. A. Epiphallus, lateral view. B. Epiphallus, axial view. C. Epiphallus, dorsal view. D. Phallic complex, lateral view. E. Endophallic plate. F. Ventral view of distal tip of ectophallus, to show medial ventral invagination of membrane. G. Dorsal view of tip of phallic complex. H. Axial view of phallic complex, epiphallus omitted.
Fig. 8.
Homeomastax bustum n. sp., male genitalia. A. Epiphallus, dorsal view. B. Epiphallus, axial view. C. Entire phallic complex, lateral view. D. Parasagittal section through C at the level of the ectophallic sclerite. Note that in this species and in the following two (pluvialis, acrita) the spermatophore sac joins the sclerite sac below and halfway along the ectophallic sclerite. E. Sagittal section through C. The opening of the spermatophore sac is restricted to the region shown with a dotted line. The dorsal part of the ventroapical evagination is the continuation of the floor of the distal part of the spermatophore sac. All three species have a distinct ventroapical evagination and a medial apodeme inserts at its base. F. Endophallic plate, ventral view. G. Dorsal view of tip of phallic complex. H. Endophallic plate, axial view, with sclerite sac, spermatophore sac, ejaculatory duct and ejaculatory sac. Abbreviations: ses, subepiphallic sac; scls, sclerite sac.
Fig. 9.
Homeomastax pluvialis n. sp., male genitalia. A. Epiphallus, dorsal view. B. Epiphallus, axial view. C. Entire phallic complex, lateral view. D. Parasagittal section through C at the level of the ectophallic sclerite. E. Sagittal section through C. F. Endophallic plate. G. Dorsal view of tip of phallic complex.
Fig. 10.
Homeomastax acrita n. sp., male genitalia. A. Epiphallus, dorsal view. B. Epiphallus, axial view. C. Entire phallic complex, lateral view. The arrow indicates the densely sclerotized lateral plaque of the apical shield. D. Parasagittal section through C at the level of the ectophallic sclerite. The arrow indicates the sclerifications in the superior wall of the dorsal fold. E. Sagittal section through C. Arrow indicates the point at which the sclerite sac and the spermatophore sac open into the genital trough. F. Endophallic plate. G. Dorsal view of tip of phallic complex. Arrows indicate the sclerified plaques in the dorsal fold and on the apical shield. Scale bar, 1 mm.
Fig. 11.
Homeomastax quinteroi n. sp., male genitalia. A. Phallic complex, dorsal view. B. Phallic complex, axial view. C. Endophallic plate, ventral view. D. Phallic complex, lateral view.
Fig. 12.
Homeomastax bouvieri (Bolívar), male genitalia. A. Phallic complex, dorsal aspect, including epiphallus. B. The same, lateral aspect; pigmented spot on apical shield not shown. C. Phallic complex, ventral aspect, membrane cut away to show endophallic plate.
Fig. 13.
Homeomastax dereixi (Descamps), male genitalia. A. Phallic complex, ventral aspect, to show endophallic plate and its insertion into apical shields. B. Phallic complex, dorsal view, including epiphallus. C. Phallic complex, lateral view. D. Phallic complex, axial view.
Fig. 14.
H. jeanninae (Descamps), male genitalia. A. Phallic complex, lateral view. B. Phallic complex, axial view. C. Phallic complex, dorsal view, including epiphallus.
Fig. 15.
Homeomastax brachyptera n. sp., male genitalia. A. Epiphallus, axial view. B, C. Epiphallus, dorsal view, two different individuals. D. Phallic complex, lateral view. E. Phallic complex from underside after removal of ventral part of ectophallic membrane, to show endophallic plate and its insertion on the apical shields. F. Apex of phallic complex, ventral, to show longitudinal folds in place of ventroapical evagination. G. Apex of phallic complex, dorsal view. Fine dashed lines indicate the position of the posterior edge of the epiphallus.
Fig. 16.
Homeomastax kressi (Rehn & Rehn), male genitalia. A. Epiphallus, axial view. B. Epiphallus, dorsal view. C. Phallic complex including epiphallus, lateral view. D. Parasagittal view of phallic complex, to show relations of endophallic plate, ectophallic sclerites, and sclerites of dorsal fold. E. Sagittal view of phallic complex, showing endophallic and ectophallic sacs. F. Endophallic plate, ventral view. G. Dorsal fold, dorsal view. Cross-hatched areas are sclerites. H. Apex of phallic complex, dorsal view. Cross-hatched areas are apical shield.
Fig. 17.
H. hondurensis (Rehn & Rehn), male genitalia. A. Phallic complex, lateral view. B. Parasagittal section to show relations of endophallic plate and ectophallic sclerite. C. Sagittal section to show endophallic sacs, ventroapical evagination and its apodeme. D. Phallic complex, frontal view. E. Phallic complex, dorsal view. F. Endophallic plate, ventral view.
Fig. 18.
Homeomastax cerciata (Hebard), male genitalia. A. Phallic complex including epiphallus, lateral view. Long thin arrow indicates the minutely spiny border of the dorsal fold (see also D); broad arrow indicates ventroapical evagination; short thin arrow indicates the semisclerotized floor of ectophallic capsule. B. Parasagittal section to show relations of endophallic plate and apical shield, and the sclerite (scl.s.), dorsal fold (d.f.) and subepiphallic (sep.s) sacs. C. Sagittal view to show endophallic sacs and sclerite sac. Broad arrow indicates genital aperture, thin arrow the finger-like posterior processes of the spermatophore sac (see also G). D. Dorsal view of apical part of phallus. Thin arrow indicates the minutely spiny border to the dorsal lobe. E. Epiphallus, dorsal view. F. Epiphallus, axial view. G. Axial view of ectophallic sclerites, processes of spermatophore sac (thin arrow) and upper part of ventroapical evagination. H. Endophallic plate, ventral view.
Fig. 19.
Homeomastax tenoriensis n. sp., male genitalia. A. Epiphallus, axial view. B. Epiphallus, dorsal view. C. Endophallic plate, ventral view. D. Apex of phallic complex, dorsal view. Densely cross-hatched areas are apical shields, lightly cross-hatched areas are sclerites of dorsal fold. E. Phallic complex including epiphallus, lateral view. F. Parasagittal view of phallic complex, to show relations of endophallic plate, ectophallic sclerites, and sclerites of dorsal fold. G. Sagittal view of phallic complex, showing endophallic and ectophallic sacs and medial apodeme. Arrows in D, E and G indicate the lobes of the ventroapical evagination.
Fig. 20.
Homeomastax silvicola (Rehn & Rehn), male genitalia. A. Epiphallus, dorsal view. B. Epiphallus, axial view. C. Detail of epiphallus, dorsal view, from another individual, showing different shape of sensory area at base of lophus. D. Phallic complex, including epiphallus, lateral view. E. Parasagittal view of phallic complex, to show relations of endophallic plate, ectophallic sclerites, and sclerites of dorsal fold; s-ep.s, sub-epiphallic sac; scl.s, sclerite sac. F. Sagittal view of phallic complex, showing endophallic and ectophallic sacs. G. Endophallic plate, ventral view. H. Endophallic plate, axial view. J. Apex of phallic complex, dorsal view. In D and J the thin arrows indicate the barb on the crest of the ectophallic sclerite; the thick arrows indicate the claw-like ending of the ectophallic sclerite.
Fig. 21.
Homeomastax robertsi (Descamps), male genitalia. A. Entire phallic complex, lateral view. B. apex of same, dorsal view; d.l., dorsal lobe; a.s., apical shield; g.a. genital aperture; v.e., ventroapical evagination. C. Epiphallus, axial view. D. Epiphallus, dorsal view. E. Endophallic plate, ventral view. F. Entire phallic complex, lateral view, with fully formed spermatophore in place (broad arrow). Note the retraction of the ventral membrane (thin arrow). G. As F, but after maceration and removal of the spermatophore. Note the dorsal distension of the spermatophore sac, compared to the situation in A, and the folds of retraction of the dorsal lobe (arrow). H. As G, perspective view from above and to the left. Note spread ectophallic sclerites, the circular opening of the genital aperture (broad arrow), and the folds where the ectophallic membrane and the dorsal fold are retracted (thin arrows).
Fig. 22.
Homeomastax strigla n. sp., male genitalia. A. Phallic complex without epiphallus, dorsal view. B. Entire phallic complex, lateral view. C. Epiphallus, axial view. D. Epiphallus, dorsal view. E. Endophallic plate, ventral view. F, G. Ectophallic sclerites from a completely macerated preparation, in axial and dorsal view.
Fig. 23.
Distribution map of surda species-group. The two records of H. bustum marked with question marks correspond to the specimens erroneously described as H. dentata by Rehn & Rehn (1934); see text under the latter species for discussion.
Table 1.
Dimensions and morphometric ratios of Homeomastax surda (Burr).
Table 2.
Dimensions and morphometric ratios of Homeomastax carrikeri (Hebard).
Table 3.
Dimensions and morphometric ratios of Homeomastax veraguae (Hebard).
Table 4.
Dimensions and morphometric ratios of Homeomastax bustum n. sp.
Table 5.
Dimensions and morphometric ratios of Homeomastax pluvialis n. sp.
Table 6.
Dimensions and morphometric ratios of Homeomastax acrita n. sp.
Table 7.
Dimensions and morphometric ratios of Homeomastax quinteroi n. sp.
Table 8.
Dimensions and morphometric ratios of Homeomastax bouvieri (Bolívar).
Table 9.
Dimensions and morphometric ratios of Homeomastax dereixi (Descamps).
Table 10.
Dimensions and morphometric ratios of Homeomastax jeanninae (Descamps).
Table 11.
Dimensions and morphometric ratios of Homeomastax brachyptera n. sp.
Table 12.
Dimensions and morphometric ratios of Homeomastax kressi (Rehn & Rehn).
Table 13.
Dimensions and morphometric ratios of Homeomastax hondurensis (Rehn & Rehn).
Table 14.
Dimensions and morphometric ratios of Homeomastax cerciata (Hebard).
Table 15.
Dimensions and morphometric ratios of Homeomastax tenoriensis n. sp.
Table 16.
Dimensions and morphometric ratios of Homeomastax silvicola (Rehn & Rehn).
Table 17.
Dimensions and morphometric ratios of Homeomastax robertsi (Descamps).
Table 18.
Dimensions and morphometric ratios of Homeomastax strigla n. sp.
Table 19.
Dimensions and morphometric ratios of Homeomastax dentata (Saussure).
Table 20.
Dimensions and morphometric ratios of Homeomastax richteri (Descamps).
Table 21.
Dimensions and morphometric ratios of Homeomastax annulipes (Descamps).
Table 22.
Dimensions and morphometric ratios of Homeomastax sp. indet.
Table 23.
Comparison of dimensions and morphometric ratios for all species of Homeomastax. Figures in bold type are means, those in normal type are derived from single individuals. For more details, refer to individual species descriptions and preceding Tables.
Table 23.
(Continued).
