Ceraeochrysa intacta ( Navás, 1912 ), Navas, 1912

Tauber, Catherine A. & Flint, Oliver S., 2010, Resolution of some taxonomic and nomenclatural issues in a recent revision of Ceraeochrysa (Neuroptera: Chrysopidae), Zootaxa 2565, pp. 55-67 : 61-65

Ceraeochrysa intacta ( Navás, 1912) , genus incertae sedis

Chrysopa intacta Navás, 1912: 199 [Original syntype reported to have been retained in Navás collection, probably destroyed; Neotype, CNC; designated by Garland (1985a: 137)]. Junior subjective synonym of Cer. placita by Garland (1985a: 137). Here, recognized as a valid species. New status.

Ceraeochrysa intacta (Navás) . New combination*. We consider the genus to be uncertain and we refer to the species as Ceraeochrysa intacta (Navás) , genus incertae sedis.

Chrysopa forreri Navás, 1913 –14 [1914]: 97 [Syntype, BMNH]. Junior subjective synonym of Cer. intacta . New synonymy.

Ceraeochrysa chiricahuae Freitas and Penny , in Freitas et al. 2009: 594 [Holotype, CAS]. Junior subjective synonym of Cer. intacta . New synonymy.

* Published information concerning Cer. intacta under the name placita / placitus (as Chrysopa , Ceraeochrysa , Chrysopodes (Neosuarius) , Oviedus ) is listed by Garland & Kevan (2007: 59); other references include Tauber & de León (2001, as Cer. placita ); Tauber (2010: 12, as Chrysopodes placitus ).

Below we discuss three issues. The first concerns the name of the species that Freitas and Penny (in Freitas et al. 2009) recently described as distinct from Cer. placita and the identity of its synonyms; the second concerns morphological differences between that species and Cer. placita ; and the third addresses the generic affiliation of the two species.

Species name and synonyms. Chrysopa intacta Navás , Chrysopa forreri Navás , and Ceraeochrysa chiricahuae Freitas and Penny are here recognized as subjective synonyms. The first two names are former synonyms of Cer. placita . The species-name with precedence is C. intacta Navás.

Types: Navás stated that he retained the type of C. intacta (collected in Toronto, Canada) in his personal collection. However, it has been missing for a long time. It is not in the Natural History Museum of Barcelona (MZB) ( Garland 1985a: 137; Monserrat 1985: 237), nor was it found in the MNHN (Legrand et al. 2008). Garland (1985a: 137) designated a neotype (male), which resides in the Canadian National Collection (CNC); it was collected from Kazabazua, Québec, Canada ( Fig. 6 ), and he recognized the species as a junior subjective synonym of Cer. placita . In their revision, Freitas et al. (2009) did not indicate that they had examined the specimen. Our (CAT) examination indicates that Garland’s neotype is conspecific with Cer. chiricahuae of Freitas and Penny (in Freitas et al. 2009). It has red markings on the gena and clypeus, a pale, unmarked vertex, a slight rosy tinge sublaterally on the pronotum, sinuate branches of the radial sector, and slightly enlarged costal cells ( Figs. 4, 5 , 7 here) – all features that Freitas and Penny used to characterize Cer. chiricahuae . In addition, the genitalia closely resemble those illustrated by Freitas and Penny for Cer. chiricahuae .

Chrysopa forreri Navás was described from Mexico City, and the syntype is in the BMNH. Freitas et al. (2009) did not list the syntype among the specimens that they had examined; our (CAT) examination of the specimen (abdomen missing, sex unknown) indicates that it is conspecific with the C. intacta neotype. It shares all the external features listed above for Cer. chiricahuae and the C. intacta neotype.

In addition to the C. forreri type from Mexico City, we (CAT) have seen a number of C. intacta specimens from Mexico. There is a series of thirteen specimens from Durango [in the San Diego County Museum of Natural History (SDMNH)] and one specimen from Chihuahua, Mexico [Royal Ontario Museum, Ottawa (ROM)]. There also is a record from Michoacan, Mexico ( Oswald et al. 2002: 578). Thus, the C. forreri type currently represents the southern-most record for Cer. intacta .

Conclusion: We conclude that Ceraeochrysa intacta (Navás) is the valid name for the species that Freitas and Penny (in Freitas et al. 2009) differentiated from Cer. placita . It has two junior subjective synonyms – C. forreri and Cer. chiricahuae .

Genitalic differences between Ceraeochrysa placita and Cer. intacta females.

Background & discussion: Freitas and Penny (in Freitas et al. 2009) described genital differences between the males of Cer. placita and the species they referred to as Cer. chiricahuae (= Cer. intacta here)— specifically, they stated that the Cer. placita gonocornua are “much longer” than those of Cer. intacta . Indeed, the gonocornua of the two known male Cer. placita specimens (the MCZ syntypes) appear longer than those of Cer. intacta males; they are also more robust basally and more closely aligned mesally than those of Cer. intacta .

Freitas et al. did not treat the females of the two species in their revision. Our examination of numerous female Cer. intacta specimens and four cleared and dissected Cer. placita females [two paralectotypes, one of two non-types from New Mexico [National Museum of Natural History, Washington, DC (USNM)], and another non-type from Oregon (MCZ)] showed significant differences between the females of the two species. First, among the female Cer. placita specimens that are cleared, all have smaller spermathecal invaginations than do the Cer. intacta females that we examined (compare Figs 8 and 9 here). Second, on Cer. intacta females, the lobe at the tip of the seventh sternite (first discovered and considered to be the praegenitale by Garland 1982: 254, Fig. 116) is elongate and has well defined sides; it protrudes beyond the tip of sternite VII ( Fig. 10 b). In contrast, the Cer. placita praegenitale is shorter and compressed, with folded sides; it protrudes very little, if at all, from beneath the seventh sternite ( Fig. 10 a).

It is noteworthy that the Cer. intaca praegenitale can express two types of asymmetry. Garland (1982, 1985b; Garland & Kevan [2007, as Chrysopodes (Neosuarius) placita ] reported that the Cer. intacta praegenitale bears, asymmetrically, a single stiff, robust seta at its tip; indeed, we also found the seta, but only on some, not all specimens. However, we noted that on some specimens the tip of the praegenitale can be asymmetrically expanded on one side and not the other. The significance of the two forms of asymmetry is unknown.

Conclusion: Females of Cer. placita and Cer. intacta express consistent and taxonomically useful, genitalic differences.

Generic assignments of Ceraeochrysa placita and Cer. intacta .

Background: Freitas et al. 2009: 594 reversed Tauber’s (2003) transfer of Cer. placita (now = Cer. intacta ) from Ceraeochrysa to Chrysopodes . Tauber’s transfer of the species to Chrysopodes was made largely on the basis of a long series of features on the intacta larvae that are more typical of Chrysopodes than Ceraeochrysa . Freitas et al. (2009) made the change in generic assignment because Cer. intacta (= Cer. chiricahuae of Freitas and Penny) and Cer. placita males have two major genitalic features that are typically expressed by Ceraeochrysa , and not by Chrysopodes : an elongate, tubular gonapsis and elongate gonocornua on the gonarcus.

Discussion: We agree with Freitas et al. (2009) that the presence of Ceraeochrysa -like gonapsides and gonocornua on Cer. intacta and Cer. placita are inconsistent with the species’ placement in Chrysopodes . Nevertheless, we continue to question the assignment of the two species to Ceraeochrysa for two reasons:

(i) The female genitalia of both Cer. intacta and Cer. placita ( Figs. 8, 9 here) differ markedly from those of all other known Ceraeochrysa females. Specifically, Ceraeochrysa females have characteristically elongate, U-shaped spermathecae that open to the bursa copulatrix via an elongate slit; they also have deep invaginations (see illustrations in Adams & Penny 1985; Brooks & Barnard 1990; Freitas & Penny 2001; Freitas et el. 2009; Tauber 2003). In contrast, females of both Cer. intacta and Cer. placita have pillboxshaped spermathecae that connect to the bursa via the vellum and a short bursal duct; they also have shallow invaginations [described and figured by Garland (1982, as Oviedus placitus , unpublished combination; Tauber 2003, as Chrysopodes placita )]. None of these features are typical of either Ceraeochrysa or Chrysopodes . Moreover, females of both Cer. intacta ( Garland 1982: 254, Fig. 116; Garland 1985b: 741) and Cer. placita (see Fig 10 a, b here) have a lobe (praegenitale) at the tip of the seventh sternite that has not been reported for any other Ceraeochrysa or Chrysopodes species.

(ii) Despite the statement by Freitas et al. (2009: 505) to the contrary, an extensive suite of diagnostic larval features has been shown to distinguish Ceraeochrysa from other neotropical chrysopid genera ( Tauber et al. 1998, 2000; Tauber 2003). The characters are from both Semaphoront A (first instar: 12 characters) and Semaphoront B (second and third instars: 18 characters each); they were scored for the larvae of twelve Ceraeochrysa species, including Cer. intacta (as " placita ") and seven Chrysopodes species ( Tauber 2003). The results of Tauber’s (2003) study demonstrated numerous, very striking differences between the Cer. intacta larvae and those of other Ceraeochrysa species. The larvae showed strong [albeit, incomplete] similarity with those of the Chrysopodes species.

Conclusion: The generic assignments of Cer. intacta and Cer. placita continue to present a problem because their male, female and larval characters appear to provide contradictory information. Specifically, the male genitalia share more features with Ceraeochrysa than with Chrysopodes ; the female genitalia are not consistent with those of either Ceraeochrysa or Chrysopodes ; and, the larval characters (Cer. intacta ) are much closer to Chrysopodes than to Ceraeochrysa .

Given the ambiguities, we propose that it is preferable to admit uncertainty concerning the two species’ generic assignments than it is to place the species in an inappropriate genus. Thus, we recommend retaining the species in Ceraeochrysa temporarily, recognizing that Ceraeochrysa is paraphyletic, and referring to the species as Ceraeochrysa placita (Banks) , genus incertae sedis and Ceraeochrysa intacta (Navás) , genus incertae sedis until further studies are conducted. This recommendation highlights the need for broadly based research that ultimately promotes nomenclatural stability.

Freitas et al. (2009: 597) claim that larval features should not be used in assigning chrysopid species to genera because the larvae of too many species are unknown; they stated that doing so “… raises doubts about dozens of other species and great nomenclatorial instability.” We disagree strongly with this stance, and we refer readers to the following literature that substantiates the taxonomic value of chrysopid larval characters ( Principi 1948–49; Díaz-Aranda & Monserrat 1995; Tsukaguchi 1995; Díaz Aranda et al. 2001; Tauber 2003; Tauber et al. 2006). Rather than ignoring the larval characters, we encourage neuropterists to obtain and characterize the larvae of previously unstudied taxa.

Figures

FIGURE 10. Praegenitale of (a) Ceraeochrysa placita (Navás), genus incertae sedis and (b) Ceraeochrysa intacta (Navás), genus incertae sedis. The scale applies to both illustrations. Abbreviations: prae, praegenitale; sg, subgenitale; S 7, seventh abdominal sternite.

FIGURES 6 – 9. Ceraeochrysa intacta. 6, Labels, neotype (CNC). 7, Left wings, neotype (CNC). 8, Spermatheca, Contra Costa County, California [Essig Museum, University of California, Berkeley (UCB)]. 9, Ceraeochrysa placita, Spermatheca, paratype (MCZ).

FIGURES 4 – 5. Chrysopa intacta, neotype (CNC). 4, Head, frontolateral. 5, Head, thorax, dorsolateral.