Research ArticleNothochrysinae (Neuroptera Chrysopidae)New Larval Description and Generic Synonymy witha Consideration of Generic Relationships

Catherine A Tauber12

1 Department of Entomology Comstock Hall Cornell University Ithaca NY 14853-2601 USA2Department of Entomology and Nematology University of California Davis CA 95616 USA

Correspondence should be addressed to Catherine A Tauber cat6cornelledu

Received 13 February 2014 Revised 5 April 2014 Accepted 6 April 2014 Published 11 June 2014

Academic Editor Jacques Hubert Charles Delabie

Copyright copy 2014 Catherine A TauberThis is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

Semaphorant B of Kimochrysa africana (Kimmins) expresses all of the larval synapomorphies that characterize the subfamilyNothochrysinae Except for its head markings the larva appears identical to that of Hypochrysa elegans (Burmeister) Based onconsideration of both larval and adult similarities Kimochrysa (Tjeder) is designated to be a subjective synonym of HypochrysaHagen (New Synonymy)Themorphological basis for a previously proposed generic subdivision of Nothochrysinae is evaluated theresults indicate that the subfamily can be organized into two generic groupings each with distinct suites of shared adult charactersAs yet apomorphic support is not forthcoming from adult characters and unfortunately larvae are known from only a few generain the subfamily

1 Introduction

Chrysopid taxonomists generally agree that the subfamilyNothochrysinae is an archaic probably monophyletic group-ing [1ndash7] Nevertheless synapomorphic adult features for thesubfamily have been elusive and proposals regarding themonophyly of the subfamily largely rested on the retentionof presumed plesiomorphic character states mostly in wingvenation

In contrast recent investigations have identified severallarval features (listed in a later section) that may lendapomorphic support for the monophyly of Nothochrysinae[7ndash10] These studies also indicate that larvae within thesubfamily express a range of variation in their overall bodyformmdashfrom naked to light debris carrying

To date larvae are described for only three of the ninegenera of Nothochrysinae thus the taxonomic breadth of therecently recognized morphological support for the subfamilyis limited and the range and the phylogenetic value of larvalvariation among genera remain unknown In the mid 1800s

Brauer [11] provided the first description of a larva fromthe Nothochrysinae his article illustrated and described themonotypic European Hypochrysa Hagen (Semaphorant Bmdashsecond or third instar as Hypochrysa nobilis Heyd) Morerecently modern descriptions of the first and third instarsof this species appeared [8 as Hypochrysa elegans Esben-Petersen] All instars of the lone North American and the twoEuropean species of Nothochrysa McLachlan are described(see [7 8 12]) Finally the first instars of Dictyochrysa fulvaEsben-Petersen were described and compared with those ofNothochrysa [13]

In 2010 Duelli et al [14] published images and biologicalnotes on the larvae of Kimochrysa africana (Kimmins)Theseauthors made the specimens available for morphologicalstudy and description Here I describe and compare the Kafricana larvae with those known from other Nothochrysi-nae As a result of the comparison the genus KimochrysaTjeder is shown to be synonymous with Hypochrysa andquestions arise concerning the currently held generic group-ings of Nothochrysinae These questions are addressed

Hindawi Publishing CorporationPsycheVolume 2014 Article ID 839261 10 pageshttpdxdoiorg1011552014839261

2 Psyche

2 Materials and Methods

The specimenswere collected in the Republic of SouthAfricaHoek-se-Berg Pass nr Bushmans Kloof 32∘07101584004910158401015840 S19∘10101584029710158401015840 E 650m 2-X-2004 (see [14]) Unfortunately anearly shipment of larvae was lost in the mail the secondshipment contained second instars (119899 = 3) preserved inalcohol Upon arrival the specimens were photographedand the external gross features were described One of thespecimens was cleared in KOH and transferred to glycerinefor examination of fine structures and setation Two speci-mens are now returned to P Duelli Swiss Federal ResearchInstitute WSL BirmensdorfZurich Switzerland the clearedspecimen is in the Tauber Research Collection

Morphological terminology and chaetotaxy followed theusage of Rousset [15] Tsukaguchi [16] Tauber et al [17]and Monserrat and Dıaz-Aranda [8] The larval stage of theChrysopidae typically includes three instars the first of which(Semaphorant A) is markedly different from the latter two(Semaphorant B) which resemble each other very closelyexcept for size and small differences in chaetotaxy Thus itis appropriate to compare the second instars described herewith third instars of other species

3 Second Instar Kimochrysa africana

31 Diagnosis The K africana larvae express the three fea-tures previously identified as potential synapomorphies forNothochrysinae [7] including (i) antenna terminal segmentis short (simten times shorter than remainder of antenna) (ii)antenna terminus has a group of small apical setae not anelongate seta (iii) labial palpus terminal segment has morethan three lateral sensilla

As Duelli et al [14] noted the K africana larvae closelyresemble those of the nothochrysine H elegans in theirelongated naked bodies and their bright green colorationThey also share the following morphological features (i)head primary setae are blunt (ii) thorax lateral tubercles(LTs) are absent (iii) abdomen LTs are absent (iv) abdomenlaterodorsal tubercles are absent or very small and with onlyone seta and (v) types of setae thoracic notal setae are shortblunt to slightly clavate and abdominal (A1ndashA6) submediansetae are short blunt to clavate without hooks

Themost notable difference between the larvae of the twospecies is that K africana lacks the dark elongated medianhead marking of H elegans

32 Description

321 Body (Figures 1(a) and 1(b)) Length sim59-69mm(measured in lateral view through spiracles) depth sim085ndash11mm (thickest section of abdomen) Bright green colorationof living specimens [14] faded in preserved specimens dorsalsurface largely cream-colored to tan with pronotal scleritesbrown with pair of broad reddish brown vertical bandsalong lateral margins extending from cervix to tip of A9 Allsetae short smooth pale and of two types ldquobluntclavaterdquowith blunt or slightly enlarged tip usually erect and straight(primary cranial setae dorsal thoracic and abdominal setae

(submedian setae SMS)) ldquosimplerdquo with acute tip usuallyerect and slightly curved (setae on cephalic appendages somesmall usually secondary cranial setae some very small setaeon thoracic notum setae on legs posterior part of A9 A10and ventral setae)

322 Head (Figures 1(d)ndash1(f) 2(a) and 2(b)) Dorsumcream-colored with light brown to brown markings as in Figures1(d) (dorsal) 1(e) (ventral) and 1(f) (lateral) eyes with stem-mata clear surrounding integument dark brown craniumtapered posteriorly and roughly triangular with roundedposterior (dorsal view) width (across eyes) sim060ndash064mmlength (dorsum) sim052ndash054mm and depth (midregion totop of eye) sim013mm base fully exposed Anterior margin oflabrum protruding slightly straight anteriorly and roundedlaterally All dorsal primary cephalic setae present (Figures2(a) and 2(b)) blunt to slightly knobbed apically two Vxsetae present labrum with two or three pairs of setae (onemesally and two laterally) dorsum with several secondarysetae one long indistinguishable from posterior primarysetae others very short mesal to S6 Ventral primary setae(S9 S10) present S8 absent

323 Cephalic Appendages (Figures 1(d)ndash1(f) 2(a) and 2(b))Mandible long thin with length sim10-11mm width sim010mm ratio of mandible length to head width 154ndash163ratio of mandible length to head length (dorsal) 190ndash100Mandible slightly upturned distally with single acute baso-lateral seta terminus sharp with six teeth Antennal length090ndash097mm sim17-18x length of cranium width sim003-004mm (at widest part of pedicel) scape with straight sidestwo pairs of distal setae (one lateral other mesal) pedicellong about 17ndash19x length of flagellum slightly broader thanthe base of flagellum flagellum short stubby basal flagellom-ere with or without mesal seta terminus with short basolat-eral seta several very short setae Labial palp long slendersim075x length of mandible basal segment with one shortdorsal seta basally three setae distally (twomesal one lateral)middle segment long with long undivided basal subsegmentbearing sim five setae five shorter mesal subsegments with oneseta mesally elongated terminal subsegment with two longsetae distally terminal segment simone-third length of middlesegment slightly tapered distally terminus with several verysmall setae palpiger erect with relatively straight sideswith one mesal seta one lateral seta mentum with smoothplate mesal to stipes lateral to palpiger with three pairs oflong setae cardo and stipes elongate narrow longitudinallyarranged cardo behind stipes Cervix expanded laterallyventrally withdrawn from cranium dorsally with pair ofsetae dorsolaterally two pairs laterally two pairs ventrally

324Thorax (Figures 1(c) and 3(a)) Lateral tubercles absentdorsum with scattered short blunt to slightly clavate setaeprimary setae unidentified (except as noted below) Legs(Figure 1(g)) cream-colored coxae with diffuse light brownmarks anterolaterally trochanter femur with dark brownlongitudinal stripes anterodorsally posterodorsally with darkvertical stripe at tip of segment tibia with dark brown vertical

Psyche 3

025mm

(a)

(b)

(c)

(g)

(f)

(e)

(d)

1mm

025mm

1mm

05mm

05mm

025mm

Figure 1 Hypochrysa africana Kimmins second instar (Hoek-se-Berg Pass South Africa) (a) Body dorsal (b) Body lateral (c) Thoraxdorsal (d) Head dorsal (e) Head ventral (f) Head lateral (g) Legs dorsal

4 Psyche

ped

sc

S4 S10

S7

S9

co

st

pg

cm

S6

S5

(a) (b)

Vx

S1

S3

S12

lp1

lp2

lp3

lp1

lp2

S2

S11

Sen

md

md

mx

mx

025mm

fl

Figure 2 Head of Hypochrysa africana Kimmins second instar (a) Dorsal (b) Ventral Scale applies to both (a) and (b) Abbreviations cocardo cm cranial margin fl flagellum lpx labial palpus number of segment md mandible mx maxilla ped pedicel pg palpiger scscape sen sensilla stp stipes Sx primary seta number Vx Vx setae

stripe basally short diffuse light brown longitudinal marksanterobasally posterobasally tarsus claw empodium darkbrown

Prothorax (T1) with two well delineated subsegmentsseparated by transverse depression Sc1 brown elongatedextending almost to the full length of anterior subsegment

narrow anteriorly posteriorly broad mesally embeddedwithin lateral stripe delineated laterally by curved cream-colored strip Sc2 elongated narrow extending posteriorly tomargin of anterior subsegment

Mesothorax (T2) consisting of three well delineatedsubsegments separated by distinct depressions Anterior

Psyche 5

T1

T2

T3

dep

Sc1

Sc2

Sc2

Sc2

Sc3

Sc3

sp

sc

sc

sc

sp

sp

sp

S1Sc1 S2Sc1

dep

A1

A2

A7

A8 A10

A9

dep

dep

(a)

(b)

(c) (d)

dep

dep

dep

dep

dep

dep

dep

dep

Figure 3Thorax and abdomen (dorsal) ofHypochrysa africanaKimmins second instar (a)Thorax (b) First and second abdominal segments(c) Seventh and eighth abdominal segments (d) Ninth and tenth abdominal segments Abbreviations Ax number of abdominal segmentdep smooth-surfaced intrasegmental depression between subsegments S1Sc1 S2Sc1 first and second primary setae on Sc1 Sc2 and Sc3 thefirst second and third primary sclerites of each segment sc abdominal sclerite sp spiracle Tx number of thoracic segment

subsegment with pair of small setae on anterior margin(probably S1Sc1 S2Sc) but Sc1 not distinguished spiraclessimple sessile circular brown with cylindrical taperingatrium Second subsegment with pair of small sclerites (Sc2)on anterior margin each with single very small seta (S1Sc2)subsegment separated mesally from posterior subsegmentby transverse depression with pair of large bifurcatingsclerites (Sc3) laterally Sc3 with anterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of subsegment center of Sc3markedwith brownembedded in lateral stripe entirely delineated by cream-colored area

Metathorax (T3) consisting of three well delineatedsubsegments separated by transverse depressions each withpair of transversely elongated sclerites (Sc2 Sc3) laterallySc3 large with bifurcating armsmdashanterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of segment center of sclerite marked withbrown embedded in lateral stripe entirely delineated bycream-colored area

325 Abdomen (Figures 3(b)ndash3(d)) Lateral tubercles (LTs)laterodorsal tubercles (LDTs) absent spiracles (A1ndashA8) cir-cular sessile with simple cylindrical tapering atrium Each

segment (A1ndashA7) divided into three subsegments separatedby transverse depressions spiracle located laterally on dor-sum of second subsegment second third subsegments sepa-rated by pronounced depression bearing pair of clear sublat-eral sclerites A1 anterior subsegment small spindle shapedseparated from second subsegment by small depressiondorsum with single pair of SMS second third subsegmentslonger broader similar in size and setation to those of A2 A3A2ndashA8 subsegments roughly quadrangular of similar sizeextending to margin of segment separated from adjoiningsubsegments by pronounced depressions dorsum of eachsubsegment with five (four on anterior subsegment of A8)to 14 pairs of SMS A4ndashA8 pleural region (dorsal region ofsecond and third subsegments) with cluster of four to sevenshort setae A9 cylindrical with three subsegments (dorsalview) anterior subsegment without setae posterior twosubsegments sclerotized with setae mesal subsegment withsimseven pairs of dorsal setae posterior subsegment with sim12pairs of dorsal setae about half of them lateral A10 dorsumwith transverse brown band separated mesally by largearrow-shaped darker brown mark segment with simfive pairsof setae numerous microsetae All segments with few smallldquosimplerdquo or ldquodenticulaterdquo setae Venter with subsegmentation

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

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BioinformaticsAdvances in

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Signal TransductionJournal of

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Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Nucleic AcidsJournal of

Volume 2014

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

2 Psyche

2 Materials and Methods

The specimenswere collected in the Republic of SouthAfricaHoek-se-Berg Pass nr Bushmans Kloof 32∘07101584004910158401015840 S19∘10101584029710158401015840 E 650m 2-X-2004 (see [14]) Unfortunately anearly shipment of larvae was lost in the mail the secondshipment contained second instars (119899 = 3) preserved inalcohol Upon arrival the specimens were photographedand the external gross features were described One of thespecimens was cleared in KOH and transferred to glycerinefor examination of fine structures and setation Two speci-mens are now returned to P Duelli Swiss Federal ResearchInstitute WSL BirmensdorfZurich Switzerland the clearedspecimen is in the Tauber Research Collection

Morphological terminology and chaetotaxy followed theusage of Rousset [15] Tsukaguchi [16] Tauber et al [17]and Monserrat and Dıaz-Aranda [8] The larval stage of theChrysopidae typically includes three instars the first of which(Semaphorant A) is markedly different from the latter two(Semaphorant B) which resemble each other very closelyexcept for size and small differences in chaetotaxy Thus itis appropriate to compare the second instars described herewith third instars of other species

3 Second Instar Kimochrysa africana

31 Diagnosis The K africana larvae express the three fea-tures previously identified as potential synapomorphies forNothochrysinae [7] including (i) antenna terminal segmentis short (simten times shorter than remainder of antenna) (ii)antenna terminus has a group of small apical setae not anelongate seta (iii) labial palpus terminal segment has morethan three lateral sensilla

As Duelli et al [14] noted the K africana larvae closelyresemble those of the nothochrysine H elegans in theirelongated naked bodies and their bright green colorationThey also share the following morphological features (i)head primary setae are blunt (ii) thorax lateral tubercles(LTs) are absent (iii) abdomen LTs are absent (iv) abdomenlaterodorsal tubercles are absent or very small and with onlyone seta and (v) types of setae thoracic notal setae are shortblunt to slightly clavate and abdominal (A1ndashA6) submediansetae are short blunt to clavate without hooks

Themost notable difference between the larvae of the twospecies is that K africana lacks the dark elongated medianhead marking of H elegans

32 Description

321 Body (Figures 1(a) and 1(b)) Length sim59-69mm(measured in lateral view through spiracles) depth sim085ndash11mm (thickest section of abdomen) Bright green colorationof living specimens [14] faded in preserved specimens dorsalsurface largely cream-colored to tan with pronotal scleritesbrown with pair of broad reddish brown vertical bandsalong lateral margins extending from cervix to tip of A9 Allsetae short smooth pale and of two types ldquobluntclavaterdquowith blunt or slightly enlarged tip usually erect and straight(primary cranial setae dorsal thoracic and abdominal setae

(submedian setae SMS)) ldquosimplerdquo with acute tip usuallyerect and slightly curved (setae on cephalic appendages somesmall usually secondary cranial setae some very small setaeon thoracic notum setae on legs posterior part of A9 A10and ventral setae)

322 Head (Figures 1(d)ndash1(f) 2(a) and 2(b)) Dorsumcream-colored with light brown to brown markings as in Figures1(d) (dorsal) 1(e) (ventral) and 1(f) (lateral) eyes with stem-mata clear surrounding integument dark brown craniumtapered posteriorly and roughly triangular with roundedposterior (dorsal view) width (across eyes) sim060ndash064mmlength (dorsum) sim052ndash054mm and depth (midregion totop of eye) sim013mm base fully exposed Anterior margin oflabrum protruding slightly straight anteriorly and roundedlaterally All dorsal primary cephalic setae present (Figures2(a) and 2(b)) blunt to slightly knobbed apically two Vxsetae present labrum with two or three pairs of setae (onemesally and two laterally) dorsum with several secondarysetae one long indistinguishable from posterior primarysetae others very short mesal to S6 Ventral primary setae(S9 S10) present S8 absent

323 Cephalic Appendages (Figures 1(d)ndash1(f) 2(a) and 2(b))Mandible long thin with length sim10-11mm width sim010mm ratio of mandible length to head width 154ndash163ratio of mandible length to head length (dorsal) 190ndash100Mandible slightly upturned distally with single acute baso-lateral seta terminus sharp with six teeth Antennal length090ndash097mm sim17-18x length of cranium width sim003-004mm (at widest part of pedicel) scape with straight sidestwo pairs of distal setae (one lateral other mesal) pedicellong about 17ndash19x length of flagellum slightly broader thanthe base of flagellum flagellum short stubby basal flagellom-ere with or without mesal seta terminus with short basolat-eral seta several very short setae Labial palp long slendersim075x length of mandible basal segment with one shortdorsal seta basally three setae distally (twomesal one lateral)middle segment long with long undivided basal subsegmentbearing sim five setae five shorter mesal subsegments with oneseta mesally elongated terminal subsegment with two longsetae distally terminal segment simone-third length of middlesegment slightly tapered distally terminus with several verysmall setae palpiger erect with relatively straight sideswith one mesal seta one lateral seta mentum with smoothplate mesal to stipes lateral to palpiger with three pairs oflong setae cardo and stipes elongate narrow longitudinallyarranged cardo behind stipes Cervix expanded laterallyventrally withdrawn from cranium dorsally with pair ofsetae dorsolaterally two pairs laterally two pairs ventrally

324Thorax (Figures 1(c) and 3(a)) Lateral tubercles absentdorsum with scattered short blunt to slightly clavate setaeprimary setae unidentified (except as noted below) Legs(Figure 1(g)) cream-colored coxae with diffuse light brownmarks anterolaterally trochanter femur with dark brownlongitudinal stripes anterodorsally posterodorsally with darkvertical stripe at tip of segment tibia with dark brown vertical

Psyche 3

025mm

(a)

(b)

(c)

(g)

(f)

(e)

(d)

1mm

025mm

1mm

05mm

05mm

025mm

Figure 1 Hypochrysa africana Kimmins second instar (Hoek-se-Berg Pass South Africa) (a) Body dorsal (b) Body lateral (c) Thoraxdorsal (d) Head dorsal (e) Head ventral (f) Head lateral (g) Legs dorsal

4 Psyche

ped

sc

S4 S10

S7

S9

co

st

pg

cm

S6

S5

(a) (b)

Vx

S1

S3

S12

lp1

lp2

lp3

lp1

lp2

S2

S11

Sen

md

md

mx

mx

025mm

fl

Figure 2 Head of Hypochrysa africana Kimmins second instar (a) Dorsal (b) Ventral Scale applies to both (a) and (b) Abbreviations cocardo cm cranial margin fl flagellum lpx labial palpus number of segment md mandible mx maxilla ped pedicel pg palpiger scscape sen sensilla stp stipes Sx primary seta number Vx Vx setae

stripe basally short diffuse light brown longitudinal marksanterobasally posterobasally tarsus claw empodium darkbrown

Prothorax (T1) with two well delineated subsegmentsseparated by transverse depression Sc1 brown elongatedextending almost to the full length of anterior subsegment

narrow anteriorly posteriorly broad mesally embeddedwithin lateral stripe delineated laterally by curved cream-colored strip Sc2 elongated narrow extending posteriorly tomargin of anterior subsegment

Mesothorax (T2) consisting of three well delineatedsubsegments separated by distinct depressions Anterior

Psyche 5

T1

T2

T3

dep

Sc1

Sc2

Sc2

Sc2

Sc3

Sc3

sp

sc

sc

sc

sp

sp

sp

S1Sc1 S2Sc1

dep

A1

A2

A7

A8 A10

A9

dep

dep

(a)

(b)

(c) (d)

dep

dep

dep

dep

dep

dep

dep

dep

Figure 3Thorax and abdomen (dorsal) ofHypochrysa africanaKimmins second instar (a)Thorax (b) First and second abdominal segments(c) Seventh and eighth abdominal segments (d) Ninth and tenth abdominal segments Abbreviations Ax number of abdominal segmentdep smooth-surfaced intrasegmental depression between subsegments S1Sc1 S2Sc1 first and second primary setae on Sc1 Sc2 and Sc3 thefirst second and third primary sclerites of each segment sc abdominal sclerite sp spiracle Tx number of thoracic segment

subsegment with pair of small setae on anterior margin(probably S1Sc1 S2Sc) but Sc1 not distinguished spiraclessimple sessile circular brown with cylindrical taperingatrium Second subsegment with pair of small sclerites (Sc2)on anterior margin each with single very small seta (S1Sc2)subsegment separated mesally from posterior subsegmentby transverse depression with pair of large bifurcatingsclerites (Sc3) laterally Sc3 with anterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of subsegment center of Sc3markedwith brownembedded in lateral stripe entirely delineated by cream-colored area

Metathorax (T3) consisting of three well delineatedsubsegments separated by transverse depressions each withpair of transversely elongated sclerites (Sc2 Sc3) laterallySc3 large with bifurcating armsmdashanterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of segment center of sclerite marked withbrown embedded in lateral stripe entirely delineated bycream-colored area

325 Abdomen (Figures 3(b)ndash3(d)) Lateral tubercles (LTs)laterodorsal tubercles (LDTs) absent spiracles (A1ndashA8) cir-cular sessile with simple cylindrical tapering atrium Each

segment (A1ndashA7) divided into three subsegments separatedby transverse depressions spiracle located laterally on dor-sum of second subsegment second third subsegments sepa-rated by pronounced depression bearing pair of clear sublat-eral sclerites A1 anterior subsegment small spindle shapedseparated from second subsegment by small depressiondorsum with single pair of SMS second third subsegmentslonger broader similar in size and setation to those of A2 A3A2ndashA8 subsegments roughly quadrangular of similar sizeextending to margin of segment separated from adjoiningsubsegments by pronounced depressions dorsum of eachsubsegment with five (four on anterior subsegment of A8)to 14 pairs of SMS A4ndashA8 pleural region (dorsal region ofsecond and third subsegments) with cluster of four to sevenshort setae A9 cylindrical with three subsegments (dorsalview) anterior subsegment without setae posterior twosubsegments sclerotized with setae mesal subsegment withsimseven pairs of dorsal setae posterior subsegment with sim12pairs of dorsal setae about half of them lateral A10 dorsumwith transverse brown band separated mesally by largearrow-shaped darker brown mark segment with simfive pairsof setae numerous microsetae All segments with few smallldquosimplerdquo or ldquodenticulaterdquo setae Venter with subsegmentation

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

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BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

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Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 3

025mm

(a)

(b)

(c)

(g)

(f)

(e)

(d)

1mm

025mm

1mm

05mm

05mm

025mm

Figure 1 Hypochrysa africana Kimmins second instar (Hoek-se-Berg Pass South Africa) (a) Body dorsal (b) Body lateral (c) Thoraxdorsal (d) Head dorsal (e) Head ventral (f) Head lateral (g) Legs dorsal

4 Psyche

ped

sc

S4 S10

S7

S9

co

st

pg

cm

S6

S5

(a) (b)

Vx

S1

S3

S12

lp1

lp2

lp3

lp1

lp2

S2

S11

Sen

md

md

mx

mx

025mm

fl

Figure 2 Head of Hypochrysa africana Kimmins second instar (a) Dorsal (b) Ventral Scale applies to both (a) and (b) Abbreviations cocardo cm cranial margin fl flagellum lpx labial palpus number of segment md mandible mx maxilla ped pedicel pg palpiger scscape sen sensilla stp stipes Sx primary seta number Vx Vx setae

stripe basally short diffuse light brown longitudinal marksanterobasally posterobasally tarsus claw empodium darkbrown

Prothorax (T1) with two well delineated subsegmentsseparated by transverse depression Sc1 brown elongatedextending almost to the full length of anterior subsegment

narrow anteriorly posteriorly broad mesally embeddedwithin lateral stripe delineated laterally by curved cream-colored strip Sc2 elongated narrow extending posteriorly tomargin of anterior subsegment

Mesothorax (T2) consisting of three well delineatedsubsegments separated by distinct depressions Anterior

Psyche 5

T1

T2

T3

dep

Sc1

Sc2

Sc2

Sc2

Sc3

Sc3

sp

sc

sc

sc

sp

sp

sp

S1Sc1 S2Sc1

dep

A1

A2

A7

A8 A10

A9

dep

dep

(a)

(b)

(c) (d)

dep

dep

dep

dep

dep

dep

dep

dep

Figure 3Thorax and abdomen (dorsal) ofHypochrysa africanaKimmins second instar (a)Thorax (b) First and second abdominal segments(c) Seventh and eighth abdominal segments (d) Ninth and tenth abdominal segments Abbreviations Ax number of abdominal segmentdep smooth-surfaced intrasegmental depression between subsegments S1Sc1 S2Sc1 first and second primary setae on Sc1 Sc2 and Sc3 thefirst second and third primary sclerites of each segment sc abdominal sclerite sp spiracle Tx number of thoracic segment

subsegment with pair of small setae on anterior margin(probably S1Sc1 S2Sc) but Sc1 not distinguished spiraclessimple sessile circular brown with cylindrical taperingatrium Second subsegment with pair of small sclerites (Sc2)on anterior margin each with single very small seta (S1Sc2)subsegment separated mesally from posterior subsegmentby transverse depression with pair of large bifurcatingsclerites (Sc3) laterally Sc3 with anterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of subsegment center of Sc3markedwith brownembedded in lateral stripe entirely delineated by cream-colored area

Metathorax (T3) consisting of three well delineatedsubsegments separated by transverse depressions each withpair of transversely elongated sclerites (Sc2 Sc3) laterallySc3 large with bifurcating armsmdashanterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of segment center of sclerite marked withbrown embedded in lateral stripe entirely delineated bycream-colored area

325 Abdomen (Figures 3(b)ndash3(d)) Lateral tubercles (LTs)laterodorsal tubercles (LDTs) absent spiracles (A1ndashA8) cir-cular sessile with simple cylindrical tapering atrium Each

segment (A1ndashA7) divided into three subsegments separatedby transverse depressions spiracle located laterally on dor-sum of second subsegment second third subsegments sepa-rated by pronounced depression bearing pair of clear sublat-eral sclerites A1 anterior subsegment small spindle shapedseparated from second subsegment by small depressiondorsum with single pair of SMS second third subsegmentslonger broader similar in size and setation to those of A2 A3A2ndashA8 subsegments roughly quadrangular of similar sizeextending to margin of segment separated from adjoiningsubsegments by pronounced depressions dorsum of eachsubsegment with five (four on anterior subsegment of A8)to 14 pairs of SMS A4ndashA8 pleural region (dorsal region ofsecond and third subsegments) with cluster of four to sevenshort setae A9 cylindrical with three subsegments (dorsalview) anterior subsegment without setae posterior twosubsegments sclerotized with setae mesal subsegment withsimseven pairs of dorsal setae posterior subsegment with sim12pairs of dorsal setae about half of them lateral A10 dorsumwith transverse brown band separated mesally by largearrow-shaped darker brown mark segment with simfive pairsof setae numerous microsetae All segments with few smallldquosimplerdquo or ldquodenticulaterdquo setae Venter with subsegmentation

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

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BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

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Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

4 Psyche

ped

sc

S4 S10

S7

S9

co

st

pg

cm

S6

S5

(a) (b)

Vx

S1

S3

S12

lp1

lp2

lp3

lp1

lp2

S2

S11

Sen

md

md

mx

mx

025mm

fl

Figure 2 Head of Hypochrysa africana Kimmins second instar (a) Dorsal (b) Ventral Scale applies to both (a) and (b) Abbreviations cocardo cm cranial margin fl flagellum lpx labial palpus number of segment md mandible mx maxilla ped pedicel pg palpiger scscape sen sensilla stp stipes Sx primary seta number Vx Vx setae

stripe basally short diffuse light brown longitudinal marksanterobasally posterobasally tarsus claw empodium darkbrown

Prothorax (T1) with two well delineated subsegmentsseparated by transverse depression Sc1 brown elongatedextending almost to the full length of anterior subsegment

narrow anteriorly posteriorly broad mesally embeddedwithin lateral stripe delineated laterally by curved cream-colored strip Sc2 elongated narrow extending posteriorly tomargin of anterior subsegment

Mesothorax (T2) consisting of three well delineatedsubsegments separated by distinct depressions Anterior

Psyche 5

T1

T2

T3

dep

Sc1

Sc2

Sc2

Sc2

Sc3

Sc3

sp

sc

sc

sc

sp

sp

sp

S1Sc1 S2Sc1

dep

A1

A2

A7

A8 A10

A9

dep

dep

(a)

(b)

(c) (d)

dep

dep

dep

dep

dep

dep

dep

dep

Figure 3Thorax and abdomen (dorsal) ofHypochrysa africanaKimmins second instar (a)Thorax (b) First and second abdominal segments(c) Seventh and eighth abdominal segments (d) Ninth and tenth abdominal segments Abbreviations Ax number of abdominal segmentdep smooth-surfaced intrasegmental depression between subsegments S1Sc1 S2Sc1 first and second primary setae on Sc1 Sc2 and Sc3 thefirst second and third primary sclerites of each segment sc abdominal sclerite sp spiracle Tx number of thoracic segment

subsegment with pair of small setae on anterior margin(probably S1Sc1 S2Sc) but Sc1 not distinguished spiraclessimple sessile circular brown with cylindrical taperingatrium Second subsegment with pair of small sclerites (Sc2)on anterior margin each with single very small seta (S1Sc2)subsegment separated mesally from posterior subsegmentby transverse depression with pair of large bifurcatingsclerites (Sc3) laterally Sc3 with anterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of subsegment center of Sc3markedwith brownembedded in lateral stripe entirely delineated by cream-colored area

Metathorax (T3) consisting of three well delineatedsubsegments separated by transverse depressions each withpair of transversely elongated sclerites (Sc2 Sc3) laterallySc3 large with bifurcating armsmdashanterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of segment center of sclerite marked withbrown embedded in lateral stripe entirely delineated bycream-colored area

325 Abdomen (Figures 3(b)ndash3(d)) Lateral tubercles (LTs)laterodorsal tubercles (LDTs) absent spiracles (A1ndashA8) cir-cular sessile with simple cylindrical tapering atrium Each

segment (A1ndashA7) divided into three subsegments separatedby transverse depressions spiracle located laterally on dor-sum of second subsegment second third subsegments sepa-rated by pronounced depression bearing pair of clear sublat-eral sclerites A1 anterior subsegment small spindle shapedseparated from second subsegment by small depressiondorsum with single pair of SMS second third subsegmentslonger broader similar in size and setation to those of A2 A3A2ndashA8 subsegments roughly quadrangular of similar sizeextending to margin of segment separated from adjoiningsubsegments by pronounced depressions dorsum of eachsubsegment with five (four on anterior subsegment of A8)to 14 pairs of SMS A4ndashA8 pleural region (dorsal region ofsecond and third subsegments) with cluster of four to sevenshort setae A9 cylindrical with three subsegments (dorsalview) anterior subsegment without setae posterior twosubsegments sclerotized with setae mesal subsegment withsimseven pairs of dorsal setae posterior subsegment with sim12pairs of dorsal setae about half of them lateral A10 dorsumwith transverse brown band separated mesally by largearrow-shaped darker brown mark segment with simfive pairsof setae numerous microsetae All segments with few smallldquosimplerdquo or ldquodenticulaterdquo setae Venter with subsegmentation

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

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BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

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Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 5

T1

T2

T3

dep

Sc1

Sc2

Sc2

Sc2

Sc3

Sc3

sp

sc

sc

sc

sp

sp

sp

S1Sc1 S2Sc1

dep

A1

A2

A7

A8 A10

A9

dep

dep

(a)

(b)

(c) (d)

dep

dep

dep

dep

dep

dep

dep

dep

Figure 3Thorax and abdomen (dorsal) ofHypochrysa africanaKimmins second instar (a)Thorax (b) First and second abdominal segments(c) Seventh and eighth abdominal segments (d) Ninth and tenth abdominal segments Abbreviations Ax number of abdominal segmentdep smooth-surfaced intrasegmental depression between subsegments S1Sc1 S2Sc1 first and second primary setae on Sc1 Sc2 and Sc3 thefirst second and third primary sclerites of each segment sc abdominal sclerite sp spiracle Tx number of thoracic segment

subsegment with pair of small setae on anterior margin(probably S1Sc1 S2Sc) but Sc1 not distinguished spiraclessimple sessile circular brown with cylindrical taperingatrium Second subsegment with pair of small sclerites (Sc2)on anterior margin each with single very small seta (S1Sc2)subsegment separated mesally from posterior subsegmentby transverse depression with pair of large bifurcatingsclerites (Sc3) laterally Sc3 with anterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of subsegment center of Sc3markedwith brownembedded in lateral stripe entirely delineated by cream-colored area

Metathorax (T3) consisting of three well delineatedsubsegments separated by transverse depressions each withpair of transversely elongated sclerites (Sc2 Sc3) laterallySc3 large with bifurcating armsmdashanterior arm extendinganterolaterally posterior arm extending posterolaterally eachtoward edge of segment center of sclerite marked withbrown embedded in lateral stripe entirely delineated bycream-colored area

325 Abdomen (Figures 3(b)ndash3(d)) Lateral tubercles (LTs)laterodorsal tubercles (LDTs) absent spiracles (A1ndashA8) cir-cular sessile with simple cylindrical tapering atrium Each

segment (A1ndashA7) divided into three subsegments separatedby transverse depressions spiracle located laterally on dor-sum of second subsegment second third subsegments sepa-rated by pronounced depression bearing pair of clear sublat-eral sclerites A1 anterior subsegment small spindle shapedseparated from second subsegment by small depressiondorsum with single pair of SMS second third subsegmentslonger broader similar in size and setation to those of A2 A3A2ndashA8 subsegments roughly quadrangular of similar sizeextending to margin of segment separated from adjoiningsubsegments by pronounced depressions dorsum of eachsubsegment with five (four on anterior subsegment of A8)to 14 pairs of SMS A4ndashA8 pleural region (dorsal region ofsecond and third subsegments) with cluster of four to sevenshort setae A9 cylindrical with three subsegments (dorsalview) anterior subsegment without setae posterior twosubsegments sclerotized with setae mesal subsegment withsimseven pairs of dorsal setae posterior subsegment with sim12pairs of dorsal setae about half of them lateral A10 dorsumwith transverse brown band separated mesally by largearrow-shaped darker brown mark segment with simfive pairsof setae numerous microsetae All segments with few smallldquosimplerdquo or ldquodenticulaterdquo setae Venter with subsegmentation

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

6 Psyche

lightly demarcated (A1ndashA4) or without demarcation (A5ndashA9) each segment with numerous ldquosimplerdquo setae usuallylonger than dorsal setae

4 Discussion

41 Synonymy Based on Larval Characters A comparisonof the K africana and H elegans larvae (Semaphorant B)does not reveal any significant generic-level differences forcomparison see [8] Indeed the larvae of the two speciesappear to differ largely in their head markings Because oftheir similarity I consider them congeneric and herebyKimochrysa Tjeder becomes a junior (subjective) synonym ofHypochrysa Hagen (New Synonymy) The type species of thegenus is Chrysopa nobilis Schneider and the species understudy here reverts to its original name Hypochrysa africanaKimmins

Given this generic synonymy the questions now are asfollows (i) Is this synonymy which is based on larval charac-ters also supported by adult morphological characters And(ii) how does the synonymy affect current understandings ofrelationships among the genera within Nothochrysinae

42 Support from Adult Morphology Of the several adultcharacters purported to differentiate Kimochrysa andHypochrysa two tend to support the synonymy one weaklycontradicts it and three are neutral (either variable orwithout sufficient comparative information) as follows

consistent wing venation [21] spiracle of eighthabdominal segment (female)mdashopening on the eighthtergite versus on membrane (see [4] also see Table 1)

contradictory ninth tergite and ectoproct (male andfemale)mdashfused versus separate (see [1 4 21] also seeTable 1)

neutral ectoproct (male)mdashwith versus without along slender ldquoappendagerdquo (apodeme) [21] subgeni-tale (female)mdashsclerotized versus unsclerotized [21]microtholi (domelike cuticular glands) on maleabdomen present versus absent (see [4] also seeTable 1)

Below I discuss the perspectives of two sets of authorswho provided evidence for separating the genera (i) Tjeder[21] and (ii) Brooks and Barnard [2] and Brooks [4]

421 Tjederrsquos Perspective Tjeder [21] described Kimochrysaon the basis of adult specimens of three South Africanspecies His generic description noted similarities in the wingvenation of Kimochrysa and Hypochrysa but he felt that sig-nificant differences in their male terminalia required placingthem in separate genera Specifically he mentioned (i) tergite9+ectoproctmdashseparate inKimochrysa (as opposed to fused inHypochrysa both sexes) (ii) male ectoproct in Kimochrysamdashlacking a long slender ldquoappendagerdquo (probably the apodeme)

that occurs in Hypochrysa and (iii) subgenitale (female)mdashunsclerotized in Kimochrysa (as opposed to sclerotized inHypochrysa)

Of Tjederrsquos above three characters subsequent studieshave shown that the first continues to provide the strongest(albeit weak) support for the separation of Kimochrysa fromHypochrysa In males of H elegans the T9+ect is fused andin Kimochrysa impar (Tjeder) they are separate (males areundescribed for the other two Kimochrysa species) (Table 1)In females the structures are partially fused in H elegansand separate in all three species of Kimochrysa It shouldbe noted that within at least two genera of Nothochrysinae(Pimachrysa Adams (males) and Nothochrysa (females))this character is known to express interspecific variationConsequently given the small number of studied specieswithin Hypochrysa and Kimochrysa (males 119899 = 1 for eachgenus females 119899 = 4 with one showing an intermediatecondition) (Table 1) this character lends only weak supportfor separating the two genera

The second character (an elongate male ectoproct) isreported fromH elegans [1 23] but not fromany other speciesin Nothochrysinae [2 4] Thus although this characterdistinguishes H elegans males from the single species ofKimochrysa whose males are studied its phylogenetic valueat the generic-level (versus species-level) remains open

The third character (sclerotized subgenitale) also is ofquestionable value at this time Tjeder illustrated the tips ofthe subgenitale of the three Kimochrysa species However hewas not specific about what he meant by the character and hedid not include equivalent drawings of the Hypochrysa sub-genitale for comparison Subsequent drawings and descrip-tions by other authors neither provide comparative informa-tion nor mention any differences between the subgenitale ofHypochrysa and Kimochrysa [2 23] Second the characterstate of the genitale has not been reported for species inother genera of Nothochrysinae for meaningful comparisonIn sum the generic-level value of the character needs furtherevaluation

422 Brooks and Barnardrsquos Perspective In subsequentstudies Brooks and Barnard [2] and Brooks [4] retainedKimochrysarsquos distinction as a genus apparently based onthree characters First like Tjeder [21] they noted that thecondition of the ninth tergite and ectoproct differentiatedHypochrysa (fused) from Kimochrysa (whose structures theyconsidered partially fused) As stated above this charactercurrently has little informative value regarding the synonymy

The second character was the presence or absence ofmicrotholi on the male abdomen This character differsbetween the single Kimochrysa species with known males(microtholi absent) and H elegans (microtholi present)(Table 1)However the condition is highly variable among thegenera of Nothochrysinae both within and outside the groupthat contains Hypochrysa (Table 1) Thus the character doesnot provide strong support either for or against the synonymy

The third character concerns the placement of the spirac-ular opening on the female eighth abdominal segmentmdashon

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 7

Table 1 Summary of adult characters currently used for classifying Nothochrysinae All known species are included

Species nameCharacter

T9 + ectlowast Microtholilowastlowast SpiraclelowastlowastlowastMale Female

Nothochrysa groupingAsthenochrysa

viridula (Adams) + [3] minus [3 18] minus [3] minus [3]Dictyochrysa

fulva Esben-Petersendagger + [2 19] + [19] minus [19]latifascia Kimmins + [20] + [19]peterseni Kimmins + [19] + [2] +minus [19]

Hypochrysaafricana Kimminsdagger minus [2 21 22] + [2 21]elegans (Burmeister)dagger + [2 23] +minus [2 23] + [2 23] + [2 23]impar (Tjeder) minus [2 21] minus [21] minus [2] + [21]raphidiodes (Tjeder) minus [21] + [21]

Nothochrysacalifornica Banksdagger + [1] [3] + [1] capitata (Fabricius)dagger + [2] + [2] fulviceps (Stephens)dagger + [2 23] minus [23] + [2 23] + [23]indigena Needham + [24] sinica Cndashk Yang minus [25] + [25]turcica Kovanci and Canbulat + [26] minus [26] + [26]

Triplochrysapallida Kimmins + [2] minus [19] minus [2] + [2]kimminsi New + [19] minus (prob) [19] + [19]

Pamochrysa groupingLeptochrysa

prisca Adams and Penny minus [18] minus [3]Pamochrysa

stellata Tjeder minus [21] minus [2 21] minus [2] minus [21]Pimachrysa

albocostalis Adams minus [1] + (prob) [1] fusca Adams minus [1 2] minus [1 2] + [1 2] minus [1 2]grata Adams minus (prob) [1] intermedia Adams minus (prob) [1] nigra Adams +minus [1] minus (prob) [1] + [1 2]

Characters lowasttergite 9 and ectoproct fused (+) unfused (minus) partially fused (+minus) lowastlowastmicrotholi on male sternites present (+) absent (minus) lowastlowastlowastlocation ofspiracular opening on female eighth abdominal segment on membranous pleuron below T8 (+) on T8 (minus) spanning both T8 and the membrane below (+minus)male or female unknown or character state not reported () daggerSpecies with larvae described Numbers in square brackets references

the membrane versus on the tergite For all know speciesof Hypochrysa and Kimochrysa the spiracle opens on themembrane (Table 1) thus it is consistent with the newsynonymy

43 Relationships among Genera of Nothochrysinae The newsynonymization of Kimochrysa with Hypochrysa led to areview and reevaluation of the current generic groupingswithin Nothochrysinae and the characters used to supportthe groupings Previously the subfamily was proposed tocontain a derived ldquomonophyleticrdquo group of five genera (theldquoNothochrysa grouprdquo) that included Hypochrysa and that

was supported by characters presumed to be apomorphicThe four remaining genera including Kimochrysa wereconsidered less derived and lacked apomorphic support [4]The present findings favor the notion that the subfamilycontains two generic groupings that are similar but notidentical to those listed earlier Moreover the strength of theldquoapomorphicrdquo support for the groupings is questioned fortwo main reasons First ancestral states for the presumedapomorphies lack strong supporting evidence Second thecharacters exhibit variation within the generic groupings andsometimes even within genera

Below is a summary of the now tentative groupingswithinNothochrysinae followed by a discussion of the underlying

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

8 Psyche

support and a recommendation for fuller comparative stud-ies

431 Groupings of Nothochrysinae Genera Nothochrysinaeis now proposed to contain two relatively distinct groupingsof genera

(i) Nothochrysa grouping This category contains fivegenera Asthenochrysa Adams Dictyochrysa Esben-PetersenHypochrysaHagen (including Kimochrysa)Nothochrysa McLachlan and Triplochrysa KimminsLarvae have been described for three of the genera [7]

(ii) Pamochrysa grouping This category contains threegenera Leptochrysa Adams and Penny PamochrysaAdams and Pimachrysa Adams Larvae are unde-scribed

432 Supporting Characters To provide perspective andencourage stronger comparative morphological studies thethree presumed ldquoapomorphic charactersrdquo for these groupingsare discussed below

(i) Character number 1mdashninth tergite (T9) and ectoproctof both sexes fused (versus separate) The ancestral state ofthis character is not determined but an unfused conditionhas been considered plesiomorphic for Chrysopidae [2]Among three neuropteran families considered to be relatedto Chrysopidae [5 6 27 28] the T9 and ectoproct areseparate in Polystoechotidae and variable (fused or unfused)in Hemerobiidae and Osmylidae [27 29ndash32] The fusedcharacter state occurs in various taxa of all three chrysopidsubfamilies including the two that are considered basal(Nothochrysinae and Apochrysinae) (for phylogenetic rela-tionships among chrysopid subfamilies see [5 6] for thedistribution of morphological features see [2 33 34])

From the available literature a fused condition occursin males of all genera in the Nothochrysa grouping and infemales of most genera The condition is largely absent frombothmales and females of the Pamochrysa grouping (Table 1)The most parsimonious conclusion from the distribution ofthe features is that the fused T9 and ectoproct began toevolve within males of the Pamochrysa grouping but that thepresumed plesiomorphic state (separate T9 and ectoproct)was largely retained by both males and females Apparentlyfull fusion appeared in males and fusion began in femalesduring or soon after the differentiation of the Nothochrysagrouping

The variability (especially partial fusion) in the expres-sion of the trait may reflect differences in the degree ofintegumental sclerotization rather than in the actual fusingof segments It is well known that chrysopid adults becomemore sclerotized as they mature (often requiring a periodof several weeks after emergence) and that their patterns ofsclerotization vary considerably individually and with age [135ndash37]Thus caution is necessary in scoring and interpretingthis character

(ii) Character number 2mdashmale sternites with microtholibeing present (versus absent) Microtholi are not knownfrom Neuroptera other than the Chrysopidae and theirabsence is probably the basal state for the family Within

the Chrysopidae microtholi are usually but not alwaysabsent in Apochrysinae males [2 34] and their occurrence ishighly variable among the Chrysopinae and Nothochrysinae(see [2] also see Table 1) It is not clear why Brooks [4]considered this feature as an apomorphy for the Nothochrysagroup when earlier Brooks and Barnard [2] reported severalgenera in the group as being without microtholi and it wasknown that the structures occur within PimachrysaThus thefeaturersquos pattern of occurrence is not consistent with its iden-tification as an apomorphy for the subfamily Nothochrysinaeor for either of its proposed generic groupings

(iii) Character number 3mdashfemale with spiracle on eighthabdominal segment opening on membrane (versus on ter-gite) The spiracular opening is consistently on the eighthtergite in Osmylidae [25 27 29 38] and Polystoechotidae[30 32] but its placement shows significant variation withinHemerobiidae [27 31] Among the Chrysopidae the openingis on the membrane throughout the Apochrysinae andin most Chrysopinae and its placement is variable withinNothochrysinae see [4 23 35] also (Table 1) The presumedbut as yet unconfirmed plesiomorphic state for the Chrysop-idae is for the eighth abdominal spiracle to open on the tergite[4]

In Nothochrysinae a spiracular opening on the eighthfemale tergite typifies the Pamochrysa grouping howeverthe numbers of exemplarsgenus are very small (119899 =1genus) In the Nothochrysa grouping a spiracular openingon the pleural membrane appears to typify three of the fivegenera (Hypochrysa Nothochrysa and Triplochrysa) but inDictyochrysa it is variable and it is absent from the singlespecies of Asthenochrysa (Table 1) An interesting interme-diate situation occurs in Dictyochrysa peterseni Kimminsin this species the spiracular opening appears to span themembrane and the tergite [19 Figure 66] Thus althoughBrooks may be correct in his proposal [4] that the spiracularopening on the membrane is a ldquostem apomorphic characterrdquofor the ldquoNothochrysa grouprdquo it appears more likely that thecharacter evolved within the Nothochrysa grouping after itsdifferentiation

It is noteworthy that all species of the Chrysopinaesubgenus Chrysopodes (Neosuarius) Adams and Penny havetheir spiracular opening on Tergite 8 [35]mdasha presumedreversal to the pleisiomorphic state In this group where largeseries of specimens are available for comparative study thereappears to be developmental and interspecific variation inthe extent and intensity of the tergitersquos lateral sclerotizationso that in some specimens especially those that are teneralthe spiracle appears to open on an unsclerotized portionof the pleuron membrane If a similar situation occurs inthe Nothochrysinae it could present a confounding factorsimilar to that in Character number 1 above

433 Future Studies From the above it is evident that thefew adult characters currently used to explain the phylogenyof Nothochrysinae (the presumed basal chrysopid group)offer interesting but extremely limited information on theevolutionary history of the group and of the subfamily Effortsto improve this situation have been hampered in large part

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Psyche 9

by the lack of specimens (both adult and larval) for manyof the rare crucial taxa It is hoped that the discussion herehelps stimulate efforts to collect and study this ancient groupso that future studies will be based on a broad ranging set ofcharacters (from comparative adult and larval morphologymolecular studies and comparative biology) and the fullrange of taxa

Conflict of Interests

The author declares that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The author thanks Peter Duelli (Swiss Federal Research Insti-tute WSL BirmensdorfZurich Switzerland) and MervynW Mansell (University of Pretoria Pretoria South Africa)for generously providing the specimens of K africanaand Maurice J Tauber for his helpful comments onthe paper The website ldquoLacewing Digital Libraryrdquo (J DOswald chief editor) was helpful during this research[httplacewingtamueduLDLlacewingcitationhtml] Theresearch benefitted from funding by the National ScienceFoundation the USDA Competitive Grants Program theNational Geographic Society and Cornell University (toMaurice J Tauber and Catherine A Tauber) and is part ofRegional Project W-3185

References

[1] P A Adams ldquoA review of the Mesochrysinae and Nothochrysi-nae (Neuroptera Chrysopidaerdquo Bulletin of the Museum ofComparative Zoology vol 135 no 4 pp 215ndash238 1967

[2] S J Brooks and P C Barnard ldquoThe green lacewings of theworld a generic review (Neuroptera Chrysopidae)rdquo Bulletin ofthe British Museum of Natural History Entomology vol 59 pp117ndash286 1990

[3] P A Adams and N D Penny ldquoReview of the South Americangenera of Nothochrysinae (Insecta Neuroptera Chrysopidaerdquoin Current Research in Neuropterology Proceedings of the 4thInternational Symposium on Neuropterology (24ndash27 June 1991Bagneres-De-Luchon Haute-Garonne France) M Canard HAspock and M W Mansell Eds pp 35ndash41 M Canard(Privately Printed) Toulouse France 1992

[4] S J Brooks ldquoAn overview of the current status of Chrysopidae(Neuroptera) systematicsrdquo Deutsche Entomologische Zeitschriftvol 44 pp 267ndash275 1997

[5] S Winterton and S de Freitas ldquoMolecular phylogeny of thegreen lacewings (Neuroptera Chrysopidae)rdquo Australian Jour-nal of Entomology vol 45 no 3 pp 235ndash243 2006

[6] N Haruyama A Mochizuki P Duelli H Naka and MNomura ldquoGreen lacewing phylogeny based on three nucleargenes (Chrysopidae Neuroptera)rdquo Systematic Entomology vol33 no 2 pp 275ndash288 2008

[7] C A Tauber M J Tauber and G S Albuquerque ldquoDebris-carrying in larval Chrysopidae unraveling its evolutionaryhistoryrdquoAnnals of the Entomological Society of America vol 107no 2 pp 295ndash314 2014

[8] V J Monserrat and L M Dıaz-Aranda ldquoLos estadios larvariosde los crisopidos ibericos (Insecta Neuroptera Chrysopidae)nuevos elementos sobre la morfologıa larvaria aplicables a lasistematica de la familiardquo Graellsia vol 68 no 1 pp 31ndash1582012

[9] L M Diaz-Aranda and V J Monserrat ldquoAphidophagous preda-tor diagnosis key to genera of European chrysopid larvae(Neur Chrysopidae)rdquo Entomophaga vol 40 no 2 pp 169ndash1811995

[10] L M Dıaz-Aranda V J Monserrat and C A Tauber ldquoChapter43 Recognition of early stages of Chrysopidaerdquo in Lacewingsin the Crop Environment P K McEwen T R New and AE Whittington Eds Cambridge University Press CambridgeUK 2001

[11] F Brauer ldquoLarve von Hypochrysa nobilis Heydrdquo in Ver-handlungen der Kaiserlich-Koniglichen Zoologisch-BotanischenGesellschaft in Wien vol 17 pp 27ndash30 1867

[12] C A Toschi ldquoThe taxonomy life histories andmating behaviorof the green lacewings of Strawberry Canyon (NeuropteraChrysopidae)rdquo Hilgardia vol 36 no 11 pp 391ndash433 1965

[13] T R New ldquoSome early stages of Dictyochrysa Esben-Petersen(Neuroptera Chrysopidae)rdquo Neuroptera International vol 1no 3 pp 136ndash140 1981

[14] P Duelli H Holzel and M W Mansell ldquoHabitat and lar-vae of the enigmatic genus Kimochrysa Tjeder (NeuropteraChrysopidae) in South Africardquo in Proceedings of the 10thInternational Symposium on Neuropterology (22ndash25 June 2008Piran Slovenia) D Devetak S Lipovsek and A E Arnett Edspp 153ndash158 University of Maribor Maribor Slovenia 2010

[15] A Rousset Morphologie Cephalique des Larves de Planipennes(Insectes Nevropteroıdes) vol 42 of Memoires du MuseumNationale drsquoHistoire Naturelle Paris Series A Zoology Editionsdu Museum 1966

[16] S Tsukaguchi Chrysopidae of Japan (Insecta Neuroptera) STsukaguchi Aioi-cho 6-14-102 Nishinomiya-shi Hyogo 662Japan (Privately published) 1995

[17] C A Tauber M J Tauber and G S Albuquerque ldquoBerch-mansus elegans (Neuroptera Chrysopidae) larval and adultcharacteristics and new tribal affiliationrdquo European Journal ofEntomology vol 103 no 1 pp 221ndash231 2006

[18] P A Adams ldquoA new species ofHypochrysa and a new subgenusand species ofMallada (Neuroptera Chrysopidae)rdquo Pan-PacificEntomologist vol 54 no 4 pp 292ndash296 1978

[19] T R New ldquoA revision of the Australian Chrysopidae (InsectaNeuroptera)rdquo Australian Journal of Zoology SupplementarySeries vol 28 no 77 pp 1ndash143 1980

[20] D E Kimmins ldquoA revision of the genera of the Apochrysinae(Fam Chrysopidae)rdquo Annals and Magazine of Natural HistorySeries 12 vol 5 no 58 pp 929ndash944 1952

[21] B Tjeder ldquoNeuroptera-Planipennia The Lace-wings of South-ern Africa 5 Family Chrysopidaerdquo in South African AnimalLife B Hanstrom P Brinck and G Rudebec Eds vol12 pp 228ndash534 Swedish Natural Science Research CouncilStockholm Sweden 1966

[22] D E Kimmins ldquoA new African Hypochrysa (Neuroptera)rdquoAnnals and Magazine of Natural History Series 10 vol 19 no110 pp 307ndash308 1937

[23] H Aspock U Aspock and H HolzelDie Neuropteren Europasvol 2 Goecke and Evers Krefeld West Germany 1980

[24] S K Ghosh ldquoContribution to the taxonomical studies ofNeuroptera (suborder Planipennia) from eastern India III

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

10 Psyche

Family Chrysopidaerdquo Records of the Zoological Survey of Indiavol 86 pp 329ndash354 1990

[25] C Yang -K ldquoNeuropterardquo in Agricultural Insects Spiders PlantDiseases and Weeds of Xizang S Zhang Ed vol 1 pp 191ndash222Xizang Renmin Press House Xizang China 1st edition 1987

[26] B Kovanci and S Canbulat ldquoA new species of the genusNothochrysaMcLachlan 1868 from northwestern Turkey (Neu-roptera Chrysopidae)with a key towestern Palaearctic speciesrdquoAnnales de la Societe Entomologique de France vol 43 no 2 pp165ndash168 2007

[27] U Aspock and H Aspock ldquoPhylogenetic relevance of genitalsclerites of Neuropterida (Insecta Holometabola)rdquo SystematicEntomology vol 33 no 1 pp 97ndash127 2008

[28] S L Winterton N B Hardy and B M Wiegmann ldquoOn wingsof lace phylogeny and Bayesian divergence time estimates ofNeuropterida (Insecta) based on morphological and moleculardatardquo Systematic Entomology vol 35 no 3 pp 349ndash378

[29] P A Adams ldquoA new genus and species of Osmylidae (Neu-roptera) from Chile and Argentina with a discussion of Pla-nipennian genitalic homologiesrdquo Postilla vol 141 pp 1ndash11 1969

[30] F M Carpenter ldquoA revision of the Nearctic HemerobiidaeBerothidae Sisyridae Polystoechotidae and Dilaridae (Neu-roptera)rdquo Proceedings of the American Academy of Arts andSciences vol 74 no 7 pp 193ndash280 1940

[31] J DOswald ldquoRevision and cladistic analysis of theworld generaof the family Hemerobiidae (Insecta Neuroptera)rdquo Journal ofthe New York Entomological Society vol 101 no 2 pp 143ndash2991993

[32] J D Oswald ldquoRediscovery of Polystoechotes gazullai Navas(Neuroptera Polystoechotidae)rdquo Proceedings of the Entomolog-ical Society of Washington vol 100 no 3 pp 389ndash394 1998

[33] S LWinterton and S J Brooks ldquoPhylogeny of the Apochrysinegreen lacewings (Neuroptera Chrysopidae Apochrysinae)rdquoAnnals of the Entomological Society of America vol 95 no 1pp 16ndash28 2002

[34] C A Tauber G S Albuquerque and M J Tauber ldquoCharac-teristics of the Loyola Navas male (Neuroptera ChrysopidaeApochrysinae)rdquo Proceedings of the Entomological Society ofWashington vol 107 no 3 pp 543ndash547 2005

[35] C A Tauber ldquoRevision of Neosuarius a subgenus ofChrysopodes (Neuroptera Chrysopidae)rdquo ZooKeys vol44 pp 1ndash104 2010

[36] CA Tauber F Sosa andG SAlbuquerque ldquoTwo commonandproblematic leucochrysine speciesmdashLeucochrysa (Leucochrysa)varia (Schneider) and L (L) pretiosa (Banks) (NeuropteraChrysopidae) redescriptions and synonymiesrdquo Zookeys vol301 pp 57ndash101 2013

[37] C A Tauber F Sosa G S Albuquerque and M J TauberldquoAdults and larvae of two Leucochrysa (Leucochrysa) species(Neuroptera Chrysopidae) descriptions biological notes andrelationshipsrdquo Zootaxa vol 3750 no 2 pp 101ndash129 2013

[38] J D Oswald ldquoTwo new South American species of the genusKempynus Navas (Neuroptera Osmylidae Kempyninaerdquo Pro-ceedings of the Entomological Society of Washington vol 96 no2 pp 367ndash372 1994

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology