Messatoporus Cushman

Santos, Bernardo F. & Aguiar, Alexandre P., 2013, <strong> Phylogeny and revision of <em> Messatoporus </ em> Cushman (Hymenoptera, Ichneumonidae, Cryptinae), with descriptions of sixty five new species </ strong>, Zootaxa 3634 (1), pp. 1-284 : 40-44

Messatoporus Cushman

Messatoporus Cushman, 1929:8–9 . Description, figure, keyed. Type species: Mesostenus discoidalis Cresson , by original designation.

Diagnosis. All Messatoporus can be recognized by the following combination of characters: ovipositor basally cylindric, apically depressed, with ventral valve completely enclosing dorsal valve as a sheath ( Figs 279–288 ); spiracle of first abdominal tergite placed approximately on midlength ( Figs 8–9 ); transverse sulcus at base of propodeum long and shallow ( Figs 7 , 162–206 ); mandible long, strongly tapered towards apex, with ventral tooth much shorter than dorsal one or indistinct ( Figs 3–4 ); posterior area of propodeum almost always transversely wrinkled ( Figs 162–206 ); fourth tarsomeres with distinct, long mesal lobe, lateral side without distinct lobe; anterior margin of propodeum concave ( Figs 7 , 170–203 ); apical area of clypeus laterally projected as triangular lobes ( Figs 2 , 139–152 ); median dorsal carina of T1 absent ( Fig. 7 ).

Female. Fore wing 4.3–14.9 mm long. Body usually slender and shiny. HEAD. Mandible long to very long, MLW 1.34–2.65, its apex much narrower than base, MWW 0.24–0.57; dorsal tooth much longer than ventral one and usually projected upwards, ventral tooth sometimes almost indistinct. Palpi densely pilose. Malar space usually very narrow, MSM 0.15–0.47. Clypeus wide, CHW 1.75–3.09, in front view almost rectangular to distinctly triangular, CWW 1.08–6.14, almost entirely flattened to moderately convex, often basally slightly convex and apically flattened; sublateral corners slightly to distinctly projected in triangular lobes; apical margin medially straight or concave, never distinctly convex, without median teeth, often very sharp and raised. Supra-clypeal area dorsally with moderately strong, somewhat U-shaped protuberance. Antenna with 21–30 flagellomeres; flagellum usually slightly enlarged towards apex, sometimes strongly widened and flattened subapically; apical flagellomere blunt, flattened, with a cluster of short and apically flattened structures. Supra-antennal area ventrally slightly to distinctly concave, usually with weak median line or carina or with median depression. Occipital carina strong, sharp, usually uniformly curved, apically usually fading out before reaching hypostomal carina, sometimes meeting hypostomal carina but usually apically weak or faint. Temple and gena in lateral view very narrow to moderately wide.

THORAX. Dorsal margin of pronotum regular, nor swollen; epomia almost indistinct to strong (absent in M. anepomius sp. nov. and M. longitergus sp. nov.), usually ending far from dorsal margin of pronotum. Mesoscutum usually strongly convex, subcircular to ovoid, 1.06–1.46 × as long as wide, almost always shiny and with fine to almost indistinct punctures; notaulus long, reaching 0.64–0.87 of mesoscutum length, weakly to deeply impressed, posterior apex always weakly impressed and joining in U-shape; surface over sternaulus smooth or with weak and short longitudinal wrinkles. Epicnemial carina usually reaching at least 0.6 of distance to subtegular ridge (except in M. bennetti sp. nov.). Sternaulus sinuous, complete, even if weaker at posterior 0.25, with or without vertical wrinkles. Median portion of posterior transverse carina of the mesothoracic venter indistinct to moderately long. Hind margin of metanotum without teeth like projections. Transverse furrow at base of propodeum very shallow, connected to median portion of the dorsal division of metapleuron forming long transverse sulcus, 0.46–1.16 × as long as anterior area of propodeum, medially without distinct longitudinal wrinkles. Juxtacoxal carina absent or represented only by very short ridges. Pleural carina absent. Fore tibia usually weakly to distinctly swollen. All fourth tarsomeres with distinct and long mesal lobe, apically with distinct cluster of bristles; lateral side without distinct lobe.

PROPODEUM: Moderately short to moderately long, 1.02–1.44 × as long as wide, almost always shiny, laterally more pilose than medially. Anterior margin centrally concave. Spiracle usually very elongate, rarely elliptic, SWL 2.00–7.4. Longitudinal carinae absent. Anterior transverse carina complete, strong, either straight or slightly arched forwards or backwards. Posterior area of propodeum with distinct, closely spaced transverse wrinkles. Posterior transverse carina usually completely absent or distinct as faint to moderately strong sublateral crests, sometimes distinguishable as a stronger wrinkle or rarely complete.

WINGS: Hyaline. Fore wing vein 1-Rs+M with bulla placed ventrally or centrally; ramellus absent (except only suggested in M. unidentatus sp. nov.); crossvein 1cu-a arising far from 1M+Rs, basad by 0.20–0.67 of its own length; vein 2Cua 0.18–0.83 × as long as crossvein 2cu-a (usually 0.3–0.5); bulla at crossvein 2m-cu moderately short, occupying 0.14–0.32 of its length, placed medially to posteriorly; cell 1+2Rs small to medium sized, APH 0.37–1.45 (usually 0.5–0.8), usually about as long as wide, rarely distinctly taller or wider, AWH 0.65–1.69, pentagonal; crossvein 3r-m at least partly spectral or rarely not differentiated; crossveins 2r-m and 3r-m (when present) parallel, subparallel or distinctly but weakly convergent, both veins usually about the same size; vein 4-Rs distinctly shorter than vein 4-M. Hind wing vein 1-M+Cu apically strongly convex; vein Cua much shorter to much longer than crossvein cu-a, HW1C 0.68–2.25; crossvein 1r-m with bulla at ventral 0.3; veins 1-Rs and 2-Rs distinctly angled, cell R1 trapezoidal; vein Cub distinct and tubular, its apical 0.5 almost always distinctly convex, rarely slightly convex or straight; vein 2-1A reaching at least 0.7 of distance to wing margin.

METASOMA: T1 very short to very long, about 0.29–0.53 × as long as T2–8, always very slender, T1LW 2.51–5.07, apex almost as wide to distinctly wider than base, T1WW 1.18–2.37 (usually 1.40–1.90), approximately cylindric to strongly compressed, without basolateral tooth, often with dorsal median depression just posterior to spiracle, sometimes also with lateral ad posterior median depressions; spiracle of T1 near its midlength, placed on basal 0.42–0.57, slightly to distinctly prominent; median dorsal carina completely absent; Dorsolateral carina absent, but traceable as faint line dividing dorsal and lateral portions; ventrolateral carina usually completely absent or traceable as a faint line, rarely distinct, though weak. Hairs on T2–8 always associated with small punctures. T2 usually short, rarely moderately long, T2LW 0.87–2.67, usually distinctly triangular, T2WW 1.31–2.87; thyridium minutely rugulose, slightly to distinctly longer than wide, rarely about as wide as high; T2 sometimes with a small depression just posterior to thyridium. T7–8 usually slightly to distinctly longer than T5–6, in species with more slender ovipositor T7–8 about as long as T5–6. Ovipositor moderately long to very long, OST 0.60–2.50 (usually 0.60–1.35), slender to moderately stout, almost always straight, rarely slightly up or downcurved, scarcely punctate, basally cylindric, apically slightly do distinctly depressed; apex of ovipositor blunt, without nodus or notch; dorsal valve without ridges or with 3–5 weak to moderately strong ridges; ventral valve apically dilated and overlapping dorsal valve as a lobe, with 4–12 weak to distinct teeth; surface in between teeth densely microsculptured.

Males. Generally similar to the respective females; for most species, males can be recognized with the key to females provided in this work. Morphological secondary sexual differences are usually more or less uniform within Cryptini , and apply to the males of Messatoporus as follows. General body size almost always smaller than respective females; however, the largest male specimens of a given species can be larger than the smallest females. Antenna with significantly more flagellomeres than in female, each flagellomere usually shorter and wider than in females; white band of flagellum starting more apically and usually covering more articles than in females of the same species; more frequently than in females, males have some or all flagellomeres of the white band only dorsally white; apical flagellomere apically pointed, without hammer-like structures. Transverse sulcus usually slightly longer than in female. Propodeum smaller, less strongly convex, propodeal wrinkles and sublateral crests, when present, usually weaker than in female. Spiracle of propodeum usually less distinctly elongate, SWL about 2.0–4.0. Fore tibia regular, not inflated. First metasomal segment more slender, T1LW about 4.4–6.7, and less triangular, T1WW about 1.1–1.8. T2–7 much more slender than in females, metasomal sternites almost always collapsed in dead specimens; apical tergites never wider than subapical ones. All those features must be understood as already implicit at the section “Male” of each species descriptions, unless otherwise indicated. For that reason, most differences listed at those sections refer to color patterns.

Male color patterns are usually similar to females, except for some important exceptions. Whitish or yellow marks on mesosoma are often more extensive, particularly on pronotum, mesopleuron and mesosternum ( M. euryoikos , Figs 51 , 128 ), but, in M. semialbiventris sp. nov. ( Figs 93 , 117 ), the male has reduced yellow marks. Fore and mid legs usually with less conspicuous blackish marks. In some species (e.g. M. euryoikos sp. nov., Fig. 51 , 128 ; M. semiaurantigaster sp. nov., Figs 105 , 129 , M. dissidens sp. nov.), the metasoma of females is mostly or entirely orange or fuscous, whereas the male metasoma has a distinct banded pattern.

Inter-specific variation. Species of Messatoporus have a wide array of variation in both general morphology and color patterns. Most body proportions are highly variable among species, as exemplified by the wide ranges of biometric ratios in the item “Female.” Also, some characters usually employed as “generic characters” among Cryptinae are rather variable in Messatoporus , such as shape and stoutness of the epomia, propodeal spiracle shape, areolet size and relative width, relative length of hind wing veins 1Cu and 1cu-a and presence/absence of apical ridges on ovipositor dorsal valve. However, the apparent high variability suggested by the total ranges can be partially misleading – most characters are fairly constant among most species of the genus, with only some outlier species for each character.

Although the general morphology varies much between species, it is usually remarkably similar among specimens of a given species. Most of intra-specific occurs in color patterns, often associated with a large distribution range (see below). It is important to note, however, that there may be two kinds of color differences: differences in the tonality color and differences in the pattern itself.

Differences in the tonality are very common and seem to be due, at least partially, to differences in collecting methods, preserving medium, and especially in the time elapsed between collecting and mounting the specimen. Specimens mounted or dried shortly after collected are usually lighter and brighter, whereas specimens which remained years or even decades in alcohol seem usually pale or darkened. M. semiaurantigaster sp. nov. is a good example. Some specimens which were mounted some months after collected have whitish (236,223,184) and bright orange (230,159,051) spots and marks, while specimens immersed in alcohol for 4–9 years before mounted have the same spots and marks in buff (224,196,124) and dark orange (206,120,056) tones. It is difficult to settle clearly which differences in tonality are due to specimen condition and which ones are real variations, unless there are specimens for which both collecting and drying date is known. For that reason, and to enable reliable identification regardless of specimen condition, differences in color tones were almost always not considered for taxonomic purposes (i.e., for species delimitation), but are described in the Variation item of species descriptions.

Sometimes, however, differences are not observed as tonalities within a same color (e.g. dark vs. light yellow), but as more distinct pattern changes. Some examples are as follows. The U-shaped mark at the mesoscutum is often reduced or almost indistinct; sometimes it is medially interrupted ( Fig. 160 ), leaving small longitudinal dashes separated from the subcircular spot on mesoscutum, or even those dashes may be indistinct. Small lateral dashes on the mesoscutum are also sometimes present, but those are frequently reduced or indistinct in some or most specimens of the same species. The central mark of the mesopleuron can also vary considerably in size within the same species (e.g. Figs 133–137 ), and can be connected or not to a second dorsal mark on the hypoepimeron (e.g. Figs 107–108 ). The area over the sternaulus, dividing the mesopleuron and mesosternum, can be entirely black in some species and only partially black in others (e.g. Figs 81–82 ). The arc-shaped posterior mark on propodeum, particularly when narrow, can be dorsally indistinct, leaving two separate posterior marks. Orange tones on legs sometimes turn to dark brown or blackish in some specimens. All those variations, although interpreted as “real” intraspecific variation, seem to have little importance to species delimitation. Those changes are almost always continuous, and frequently found in different specimens from the same locality which are otherwise almost indistinct.

Biology. Most aspects of the biology of Messatoporus remain widely unknown. Including information on previously known hosts, the new records presented herein corroborate the notion that species of Messatoporus are specialized parasitoids of mud nesting aculeate wasps. All known records are from species of Crabronidae , Pompilidae and Eumeninae (Vespidae) . The species with the most host records, M. discoidalis , attacks six species from all the three families. This relatively low host specificity is expected for idiobiont ectoparasitoids, such as most Cryptinae (Gauld 2006) . However, according to the late Ian D. Gauld (personal communication), most Ichneumonidae , even if not strictly host-specific, should not be promptly considered as “generalists”, since they usually attack hosts in specific environments, or hosts with specific morphological traits.

Although it may be consequence of sampling bias or just coincidence, it is noteworthy that there is no record of Messatoporus attacking nests of Sceliphron Klug. In Photocryptus , by contrast, a related genus for which the host records are slightly more extensive, almost all species with known host records attack nests of Sceliphron (see Aguiar and Santos 2009 for a revision).

Sampling. Most of the specimens examined in the present work were collected with Malaise traps. In part, that may be due to the fact that Malaise traps are the most commonly used sampling tool for Hymenoptera . However, it also agrees with the findings of Aguiar and Santos (2010), suggesting that Malaise traps outperform Moericke (yellow pan) traps in collecting Messatoporus . According to those data, 29.1 Moericke trap-days are needed to sample as many female specimens as one Malaise trap-day (89 Möricke traps to match one Malaise trap in both male and female specimens).

In fact, specimens of Messatoporus are usually found in a significant proportion of Malaise trap samples, at least in tropical forests (personal observation, each sample usually corresponding to seven trap-days). That, allied to the large number of specimens of the genus usually available in collections (see Material and Methods), may indicate that Messatoporus is indeed an abundant taxon.

The efficiency of other sampling methods for Messatoporus was not tested. The label information of many of the examined specimens does not state the collecting method, but some specimens were reared directly from mud nests or from trap-nests.

Distribution and biodiversity. Messatoporus occurs nearly throughout the Americas, from southern Canada to southern Argentina. There are now records from almost all countries in continental America; the exceptions are Belize, Nicaragua, French Guyana and Chile; the lack of records in all but the latter is probably simply due to lack of sampling. There are also species occurring in Cuba, Haiti, Dominican Republic, Puerto Rico and Trinidad. Because sampling effort in the Neotropical Region has been irregular and fragmentary, it is sometimes hard to detect accurately patterns of distribution and diversity, because better sampled areas tend to produce strongly biased information. Still, some general observations can be made analyzing the available records.

Species of Messatoporus usually have a wide distribution. Messatoporus transversostriatus and M. euryoikos sp. nov., for example, are recorded from all South America, from Panama to Argentina or southern Brazil. Messatoporus arcuatus is found from northeastern Mexico to Panama, and M. discoidalis , the most widely distributed species of the genus, from southern Canada to Costa Rica, encompassing almost all USA and Mexico. Some other species appear to be more restricted, e.g. occurring only at north of South America (from Amazon Basin to Panama), or on its eastern side ( M. copiosus , Fig. 316 ). Some species, like M. andinus sp. nov., are clearly Andean, occurring only above 1800 m in Peru, Colombia and Ecuador.

There are still many species which are known just from one locality, but those species are usually also represented by only one or a few specimens. Therefore, it is difficult to distinguish if those particular species have a small geographic range or if they are just less common species which were not collected in other areas because of insufficient sampling effort. Future studies and more collecting trips at Neotropical Region are likely to mitigate this problem.

The wide distribution range of many species of Messatoporus , combined with the large species richness of the genus, means that many different species can be found in a single area. Seven species of Messatoporus were recorded just from Parque Estadual do Rio Doce, a 35,974 ha state park at southeastern Brazil which was sampled with over 4,000 Malaise trap-days. At southern Bahia, extensive sampling of several close agro-forestry properties found ten species of the genus. Although it is likely that some of those species were not permanent residents of each area, those data strongly indicate that several species of Messatoporus can occur at the same area. That raises relevant questions about niche partition, host exploitation and interspecific competition. As expected for most Cryptinae and supported by the available data (see above, “Biology”), species of Messatoporus seem not to be host-specific, but to attack a wide range of hosts. How do different species diverge on their host use, in order to keep their ecological niches at least partially distinct? For Photocryptus , a very close genus, there is at least one record of two species attacking the same nest (Aguiar & Santos 2009), suggesting competition among different species. That is probably the case with Messatoporus , but it is also clear that some niche differentiation must occur.

It is also worth noting that phylogenetically close species do not usually seem to occur in sympatry. In some cases, they do occur in distinct parapatry, with the ranges of two species adjacent or close to each other but not significantly overlapping. This seems to be the case, for example, of M. lissonotus and M. paralissonotus sp. nov.; the former species occur in most of South America, from southern Brazil to Ecuador, whereas the later is recorded from Panama. The very close species M. propodeator Kasparyan & Ruíz and M. amazonensis sp. nov. do also occur in distinct parapatry, the former extending from northeastern Mexico to Costa Rica, whereas M. amazonensis is recorded from northern South America, from Northern Colombia to Central Brazil.

The situation seems to be at least partially similar to the group of very close species including M. mesosternalis , M. semiaurantigaster sp. nov., M. orientalis sp. nov., M. occidentalis sp. nov. and M. basiflavus sp. nov. These five species have almost non-overlapping records. Messatoporus mesosternalis is recorded from Mexico to Costa Rica; M. basiflavus is known only from the type locality in Panama; M. occidentalis is widely distributed at western South America. At the eastern portion of the continent, M. orientalis is recorded from northern Argentina to southeastern Brazil, in the highlands of Espírito Santo state. Messatoporus semiaurantigaster , for its turn, is recorded mainly from the lowlands of Espírito Santo, and Bahia, with a single record in southern Brazil ( Figs 345, 349, 351 , 361 ).

Figures

FIGURES 279–288. Ovipositor tip. 279, M. discoidalis. 280, M. paradoxus*. 281, M. tibiator. 282, M. citreocephalus*. 283, M. teutonicus*. 284, M. compressicornis. 285, M. nigriscutus*. 286, M. bennetti*. 287, M. semiaurantigaster*. 288, M. titans*.

FIGURES 6–9. Terms for structures used in the present work. 6, Messatoporus copiosus, dorsal view; teg, tegula; msc, mesoscutum; msl, mesoscutum length; msw, mesoscutum width; nl, notaulus; scc, scutellar carina; ssg, scuto-scutellar groove; sct, scutellum; axc1, first axillar carina; axc2, second axillar carina; psct, postscutellum. 7, M. orientalis, first metasomal tergite; tas, tergite length anterior to spiracle; tps, tergite length posterior to spiracle; dlc, dorsolateral carina. 8, M. nigriscutus, T1, dorsal view; t1l, length of T1; t1mw, minimum width of T1; t1xw, maximum width of T1. 9, M. amazonensis, T2, dorsal view; same codes of T1.

FIGURES 162–170. Propodeum, dorsal view. 162, M. advenus*. 163, M. paradoxus*. 164, M. badius*. 165, M. albiaterartus*. 166, M. longicaudis*. 167, M. amarelus. 168, M. apertus. 169, M. convexus*. 170, M. laevilatus.

FIGURES 171–179. Propodeum, dorsal view. 171, M. paraguayensis. 172, M. pallidus*. 173, M. fulvator. 174, M. nigriscutus*. 175, M. tenuirobis*. 176, M. elektor*. 177, M. versicolor*. 178, M. maculiscus. 179, M. opacus*.

FIGURES 180–188. Propodeum, dorsal view. 180, M. concavivenus*. 181, M. teutonicus. 182, M. semialbiventris*. 183, M. orientalis*. 184, M. euryoikos. 185, M. transversostriatus. 186, M. titans*. 187, M. amplithorax. 188, M. lordos.

FIGURES 189–197. Propodeum, dorsal view. 189, M. bimaculatus. 190, M. unidentatus*. 191, M. semiaurantigaster. 192, M. occidentalis. 193, M. tibiator. 194, M. dominicanus*. 195, M. jalapa. 196, M. allomeros*. 197, M. mesonotator.

FIGURES 198–206. Propodeum, dorsal view. 198, M. maculipes. 199, M. caxiuanensis*. 200, M. amazonensis. 201, M. variegatus. 202, M. atrifoveus*. 203, M. dissidens*. 204, M. complexifemur. 205, M. copiosus. 206, M. ellipsicavus*.

FIGURES 2–4. Terms for structures used in the present work. 2, Messatoporus nigriscutus. head, frontal view; aac, apical area of the clypeus; clp, clypeus; cma, clypeus maximum width; cmi, clypeus minimum width; clh, clypeus height; sca, supra-clypeal area; saa, supra-antennal area. 3, M. maculiscus; mdl, mandible length 4, M. orientalis; mls, malar space; mbw, mandible basal width; gen, gena.

FIGURES 139–152. Head, frontal view. 139, M. nigriscutus*. 140, M. transversostriatus. 141, M. maculiscus. 142, M. laevilatus. 143, M. elektor*. 144, M. versicolor*. 52. 145, M. opacus*. 146, M. semialbiventris*. 147, M. lunatus*. 148, M. bimaculatus*. 149, M. andinus*. 150, M. bennetti*. 151, M. teutonicus*. 152, M. paeneater*.

FIGURES 51–55. Lateral view. 51, M. euryoikos. 52, M. apiopharkis*. 53, M. lordos. 54, M. teutonicus*. 55, M. unidentatus*.

FIGURES 124–129. Male specimens, lateral view. 124, M. jalapa. 125, M. copiosus. 126, M. ellipsicavus. 127, M. propodeator. 128, M. euryoikos. 129, M. semiaurantigaster.

FIGURES 91–95. Lateral view. 91, M. bicolor*. 92, M. paeneater*. 93, M. semialbiventris*. 94, M. andinus*. 95, M. matucanus*.

FIGURES 115–118. 115–116: M. townesi, lateral view. 115, Lectotype, head, mesosoma, metasoma and wings. 116, Paralectotype female, mesosoma and metasoma. 117–118: Male specimens, lateral view. 117, M. semialbiventris. 118, M. andinus.

FIGURES 101–105. Lateral view. 101, M. basiflavus*. 102, M. occidentalis*. 103, M. mesosternalis. 104, M. orientalis. 105, M. semiaurantigaster.

FIGURES 153–161. Mesoscutum, dorsal view. 153, M. lissonotus. 154, M. copiosus*. 155, M. pallidus*. 156, M. semialbiventris*. 157, M. nigriangulatus*. 158, M. orientalis*. 159, M. opacus*. 160, M. bimaculatus*. 161, M. jalapa.

FIGURES 133–137. Lateral view of specimens of M. transversostriatus showing intra–specific color variation.

FIGURES 106–110. Lateral view. 106, M. amazonensis. 107, M. propodeator. 108, M. variegatus. 109, M. complexifemur*. 110, M. campanulatus*.

FIGURES 81–85. Lateral view. 81, M. atlanticus*. 82, M. caxiuanensis*. 83, M. copiosus*. 84, M. ellipsicavus. 85, M. fulvator.

FIGURES 313–320. Distribution maps for species of Messatoporus. 313, M. compressicornis. 314, M. concavivenus. 315, M. convexus. 316, M. copiosus. 317, M. covarrubiasi. 318, M. depressifrons. 319, M. dialeipsis. 320, M. discoidalis.

FIGURES 345–352. Distribution maps for species of Messatoporus. 345, M. mesosternalis. 346, M. nigriangulatus. 347, M. nigriscapus. 348, M. nigriscutus. 349, M. occidentalis. 350, M. opacus. 351, M. orientalis. 352, M. paeneater.

FIGURES 361–368. Distribution maps for species of Messatoporus. 361, M. semiaurantigaster. 362, M. subalaris. 363, M. tenuiorbis. 364, M. tenuissimus. 365, M. terebrator. 366, M. teutonicus. 367, M. tibiator. 368, M. titans.