The bee genus Caenaugochlora ( Hymenoptera , Apoidea ) and its constituent subgenera , with new species of Caenaugochlora s . str . from Ecuador

Two new species of Caenaugochlora Michener are described and fi gured from Ecuador. Caenaugochlora quichua sp. n. and C. bennetti sp. n. are both remarkable for the presence of carinate preoccipital carinae, setose compound eyes, strongly rimmed metabasitibial plate, normally pectinate inner metatibial spur, metapostnotal striae not reaching the apical margin, and the male fourth metasomal sterna with lateral projections. Th e combination of these features intermingle attributes of the presently recognized subgenera, while others are unique to the genus as a whole. A revised diagnosis is provided for the genus and its two subgenera, with Caenaugochlora s.str. accommodating C. bennetti and C. quichua, and brief comments made regarding the implications of the characters exhibited by the aforementioned species.


Introduction
Caenaugochlora was proposed by Michener (1954) for Augochlora-like bees with setose compound eyes, as a replacement for Caenohalictus Cameron which had been misapplied at the time various unrelated bees.Michener based his description on a pectinate inner metatibial spur, reduced preoccipital carina, setose compound eyes, and the fourth metasomal sternum with a medioapical notch bordered by areas with specialized setae.At that time Michener (op. cit.) included Pseudaugochlora Michener as a second subgenus but subsequent authors have considered the group as a separate genus (Eickwort 1969;Engel 2000;Michener 2007; vide Almeida 2008, for a comments regarding Pseudaugochlora).Later, Eickwort (1969) described the subgenus, Ctenaugochlora, to include a distinctive species related to Caenaugochlora.He considered Caenaugochlora proper to include those species with the fourth metasomal sternum with one or two median patches of setae on raised tubercles to either side of a medioapical depression, while Ctenaugochlora was described for a species with a densely pectinate inner metatibial spur, a metabasitibial plate only defi ned posteriorly, and the male fourth metasomal sternum with a V-shaped patch of setae bordering a slight median depression.Engel (2000) updated the number of species of both genera and diagnosed the subgenera of Caenaugochlora based on the same characters.Moure (2007) listed 15 described species for Caenaugochlora s.str., while Engel (2007Engel ( , 2009a) ) added two additional species and removed another to elsewhere in the Augochlorini.Species of Caenaugochlora s.str.described to date occur in México to Ecuador (Table 1), although unstudied and undetermined material is known from Nicaragua, Venezuela, and Trinidad.Ctenaugochlora presently includes fi ve species in Costa Rica and Panamá, with a single putative record from Bolivia (Engel and Gonçalves, in press).Nothing is known about their natural history except for the ground-nesting biology and semisocial behavior of C. costaricensis described by Michener and Kerfoot (1967).
Th e purpose of the present study is to describe two new species of Caenaugochlora s.str.from Ecuador which have diagnostic characteristics of both subgenera as currently recognized and share a peculiar structure of the fourth metasomal sternum, diff erent from that of other Caenaugochlora s.l.Both species may prove critical for illuminating the phylogenetic placement of Caenaugochlora s.l.among Augochlorini.

Methods
Material considered herein is deposited in the following institutions: Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZSP) and Snow Entomological Collection, Division of Entomology, University of Kansas Natural History Museum, Lawrence, Kansas, USA (SEMC).Quotation marks surround exact transcriptions of individual labels, while the reverse solidus (\) indicates diff erent lines on the same la- bel.Morphological terminology follows that of Eickwort (1969) and Michener (2007) except in that, following Engel (2001), we use "mesoscutum" in place of "scutum" and a revised venational nomenclature.In addition, we here use "metapostnotum" for the "basal area of the propodeum" and following Engel (2009b) we refer to the "teeth" of the pectinate spurs as "branches".Species descriptions loosely follow those of Engel (1997Engel ( , 2007)), while generic and subgeneric diagnoses and descriptions follow that of Engel (2000), with new interpretations of particular characters as revealed by the new species described herein highlighted in bold.Paratypes.3♂♂, 1♀, with same label data as that of holotype; 2♂♂, 1♀ (SEMC) and 1♂, 1♀ (MZSP) labeled, "Ecuador, Cuenca\ Azuay Prov.\III-13-1965" "L.E.Pena\ collector".

Systematics
Diagnosis.Both sexes with preoccipital ridge carinate; female metabasitibial plate well rimmed; female metapostnotum with striations weakly reaching posterior margin where they become diffi cult to discern among strongly granular integument (Fig. 3); male face almost straight above antennal alveoli, setae on this portion long and weakly branched; apical margin of metasomal sternum III not notched; apical margin of metasomal sternum IV straight, subapical margin with a distinct median patch of erect setae, length of lateral projections less than sternum length (Fig. 7).
Head punctate, punctures contiguous on upper parocular area, those punctures on frons stronger than elsewhere on face.Mesoscutum densely punctate, punctures irregular in diameter, contiguous around parapsidal lines and posteriorly, separated by less than a puncture width centrally, blending to strongly imbricate integument medioanteriorly.Mesepisternum densely punctate and rugulose.Metapostnotal striations longitudinal and partly irregular, striations covering entire surface, apically weaker and diffi cult to discern among strongly granular integument (Fig. 3).Metasomal terga lineolate with small puncures, those of tergum I smaller in diameter than those on other terga.
Integument of head and mesosoma brilliant metallic green with some cupreous refl ections.Mandible, labrum, malar space, and apical one-half of clypeus dark brown; antenna, all leg podites, tegula, and sterna light brown.Wing veins brown; membrane lightly fuscous.Terga mostly brilliant metallic green with small brown areas on median surface of metasomal terga I and II, apical marginal areas translucent brown, abruptly demarcated from remainder of terga.
Pubescence on head, mesosoma, and sterna pale yellow, on legs dark amber, on terga light amber.Setae short on terga, long and erect on sterna; metasomal sternum II and III with long setae with recurved apices.Male: as described for female except as follows: Total body length 6.0 mm; forewing length 4.8 mm; distance between compound eye notches 1.2 mm; maximum head length 1.8 mm; intertegular distance 1.1 mm.Head longer than wide; not strongly 1 2 4 6 5 1.00 mm 3 depressed (Fig. 5).Mesoscutal median line well impressed.Metapostnotum as long as mesoscutellum, slightly depressed.Lateral projection of metasomal sternum IV as long as sternum length (Fig. 7); metasomal sternum VI strongly emarginate; metasomal sternum VII and VIII as depicted in Figure 9; genital capsule as depicted in Figure 11.
Clypeus and supraclypeal area with fewer punctures than lower parocular area, with imbricate microreticulations.Mesoscutum strongly punctate, punctures separated by less than a puncture width, integument between (where perceptible) with imbricate microreticulations.Metapostnotal striae dense, almost entirely longitudinal, extending along entire surface, weaker apically and diffi cult to discern among strongly granular integument.Terga irregularly punctuate and lineolate.
Integument of head and mesosoma brilliant metallic green with some dark refl ections.Apical margin of clypeus narrowly pale amber.Coxae to femora with metallic highlights, tibiae and remaining podites dark amber.Metasoma entirely brown.
Pubescence on face long and plumose, especially above supraclypeal area.Lateral projection of metasomal sternum IV bearing four setae at apex, sternal apical margin with a central patch of setae (Fig. 7).Genital capsule as depicted in Figure 11.
Etymology.Th e specifi c epithet is a patronym honoring Mr. Daniel J. Bennett, leading student of the systematics of crabronine wasps.
Diagnosis.Both sexes with preoccipital ridge carinate; female metabasitibial plate well rimmed; female metapostnotum with striations not reaching the posterior margin; male face concave above the antennal alveolus with a stripe of short, strongly plumose setae; medioapical margin of metasomal sternum III not notched; apical margin of metasomal sternum IV straight, subapical margin with pubescence distributed as on metasomal sternum III, lateral projections, including setae, as long as sternum.

Discussion
Neither C. bennetti nor C. quichua fi t precisely the description of either subgenus in Caenaugochlora.Both species are unique among the genus by combining setose compound eyes, a carinate preoccipital ridge, and male fourth metasomal sternal projections.As previously understood, the carinate preoccipital ridge is presently only in species of Ctenaugochlora, while in Caenaugochlora s.str.the ridge is distinctly sharply angled but not carinate.On the other hand the setose compound eyes and metapostnotal ornamentation (not radiating) are typical of Caenaugochlora s.str.Th e body shape and size is reminiscent of small Caenaugochlora s.str.such as C. gemmella and C. macswaini both from southern Central America.Lastly, the presence of a broad parapenial lobe and the lateral projections of the fourth metasomal sterna are unique among Caenaugochlora s.l.Both subgenera can have median patches of setae on the fourth metasomal sternum which are frequently associated with a median notch, while the third metasomal sternum is also frequently notched and bears setae.Th e distribution of characters may suggest that they are sister to Ctenaugochlora but they may also represent a basal branch of the genus.Rather than create a third subgenus we chose to alter the delimitation of Caenaugochlora s.str.to provisionally include these new species.Together these species may prove important for understanding the phylogeny of Caenaugochlora, particularly given that the lateral projections of the male fourth metasomal sterna are similar to those found in Augochloropsis Cockerell, Augochlorodes Moure, Neocorynura joannisi (Vachal) and Th ectochlora Moure.Th ese projections in Caenaugochlora are formed by sclerotized extensions of pregradular and postgradular areas as present in some Augochloropsis (vide Eickwort 1969: 468, Figs 243, 244).For species of Th ectochlora, Augochlorodes turrifaciens Moure (Eickwort 1969: 469, Figs 248, 249) and N. joannisi (Smith-Pardo & Gonzalez 2009: 120, Fig. 13) the projections are formed principally by extension of postgradular integument, while pregradular integument does not extend along the projection.Th is trait may favor Engel's (2000) cladistic hypothesis in which Augochloropsis s.l. and Caenaugochlora s.l. were sister taxa.Modifi cations of the male sterna, especially those on the fourth sternum, are common in halictines (Michener 2007) and such lateral projections can often be convergent.Further cladistic work is needed in order to more thoroughly evaluate the precise relationship of these new species and their implications for understanding affi nities of Caenaugochlora s.l.

Figure 13 .
Figure 13.Distribution map of known Caenaugochlora s.str.from Ecuador.

Genus Caenaugochlora Michener Diagnosis.
Most species of Caenaugochlora are similar in general appearance to those of the genus Pseudaugochlora; however, the latter genus has a strong ridge on the vertex and a pointed galeal apex.From the related genus Augochloropsis, Caenaugochlora differs in the absence of a notch on the tegula, the non-lamellate pronotal dorsal ridge, and the orthogonal epistomal sulcus.Mandible with subapical tooth variously defi ned.Labral distal process narrowly triangular; basal elevation orbicular; teeth absent.Prementum not greatly elongate.Galeal apex rounded; galeal comb absent; galeal base extending to stipital base.Hypostomal ridge carinate; anterior angle rounded.Length of malar space less than basal mandibular width except in C. silvicola malar space as long as basal mandibular width.Epistomal sulcus orthogonal.Ocelli not greatly enlarged; ocellar furrow absent.Vertex not expanded or ridged behind ocelli.Preoccipital ridge sharply angled or carinate.Pronotal dorsal ridge carinate; lateral ridge rounded to carinate.Mesoscutal anterior border rounded; mesoscutal lip rounded.Tegula oval.Probasitarsal brush present; inner metatibial spur pectinate.Apex of marginal cell truncate; distal hamuli with irregular spacing pattern.Propodeal pit narrow.Male: Mandible simple.Labrum with distal process narrowly triangular as in the female; basal area notched.Antennae extending back to mesoscutellum; second fl agellomere about as long as fi rst fl agellomere.Inner metatibial spur serrate.Metasoma oval.Apical margin of metasomal sternum III variable, straight, arcuate, or notched; apical margin of metasomal sternum IV variable, lateral projections present or not; apical margins of metasomal sterna V and VI emarginate; apical margin of metasomal sternum VII with median projection; spiculum narrow; proctiger unmodifi ed.Gonobasal bridge narrow; dorsal lobes weak; basal process of gonostylus and parapenial lobe present or absent; dorsal process partly membranous; ventral surface of penis valve with prong.Compound eyes usually with long setae.Preoccipital ridge carinate (as in C. macswaini) or sharply angled (most species).Pronotal lateral angle usually produced, angle slightly acute to obtuse.Metabasitibial plate bordered anteriorly and posteriorly; inner metatibial spur pectinate.Metapostnotum with basal striae, not reaching apical margin.Male: Apical margin of metasomal sternum III variable, straight, arcuate, or notched; apical margin of metasomal sternum IV variable, straight to notched with some apical setal patches surrounding medial notch; lateral projections present or not; apical margin of metasomal sternum VIII with median projection, sometimes bilobed at apex.Volsella indented on inner margin (except in C. bennetti sp.n.).
Diagnosis.Caenaugochlora s.str. is distinguished from Ctenaugochlora by the normal pectination of the inner metatibial spur (with less than 10 long branches), frequent presence of long compound eye setae, strong anterior border of the metabasitibial plate, and metapostnotal striae not reaching to the apical margin.Description.As for the genus with the following additions: Female: Distribution.Species of Caenaugochlora s.str.occurs from México to Ecuador, including Venezuela and Trinidad.