THE WASMANN JOURNAL OF BIOLOGY

VOL. 10, No. 1 SPRING, 1952

Notes on the Biology and Host Relationships of Some Species of Nemognatha (Coleoptera: Meloidae)

E. G. LINSLEY and J. W. MAcSW AIN , D ·ivision of Entomology and Pa.1·asitology, Unive1·sity of Califontia, Be1·keley 1. CalifoTnia.

Although the meloid genus N emogn(ttha contains man:v com­ mon and widespread species, little has been recorded of the habits of the North American forms beyond the fact that several of them have been reared from the cells of bees. This has been true not only beca use few persons have investigated their habits but also results from the fact that the of the genus has been in a confused state and the determination of species has been difficult. The major portion of om previous knowl edg-e of the habits of N emognatha has been derived from the studies of Cros (1912, et seq.) upon the Old vVorld species, N. ch·rysomelina Fabricius. In general, the life histories of the species of N emogn(l,tha studied by us conform to the following basic pattern. The eggs arc laid in clusters of seYeral hunclr·ed each on the back of flower buds (plate 1, figs . 1-3) . In the plant species selected for oviposi­ tion some forms of JYemognath(t are remarkabl:v specific, others may utilize a number of genera and species in a family such as the Compositae. A few, as .\" . scut ellaTis LeConte, are associated with a great man~ · different flowers and have a wide range of host bees ( ta hi e I ). The incubation period of the eggs is usually such that the primar~ - larvae hatch soon enough to infest t he Sowers by the time the~· bloom. vVhen a bee visits the flower t he larvae attach themselves to the body hairs and are carried back to the nest. There they leave the bee and, wh en the cell has been sealed, attack and destroy the egg. Th e ~ - then feed through the fourth instar

Received for publication April 13, 195 1. [ 91 l CD N Table 1

Host relationships of some North American species of Nemognatha

N emognatha species Host Locality Authority o-j ii1 tz:j N. dttbia LeConte ·------·------··------Anthidium emm·ginatum (Say) California Davidson, 1907 llfegachile montivaga Cresson ______··· ·-- ...... California new r ecord ~ > N . huTdi Mac Swain ------····-·-·- --- 11! elissodes 1·obustio1· CockerelL ______California new record Ul ~ N . lu1·iaa LeConte .... ----· -- ·- ·· ·--··-·-· ···-· --··- -··Antho7Jho?·a occidentalis Cresson ...... Colorado Mickel, 1928 > llf egachile occidentalis Fox .. ------·- -··---·· __ ---·-· New Mexico new record z N. ltttea pallens LeConte. ______Antho co1Ja sp ..... --·· ··------·····-·------· ·- -· --· ·· ...... California new record z N. nigTipennis LeConte ...... • Dianthiditt?n Arizona Schwartz, 1903 '-< 0 llf egachile pmtti Cockerell______.. California new record q ::0 N. piezata Fabricius...... Anthophom occidentalis Cresson ...... New Mexico new record z N. scutellaTis LeConte ...... Anthophm·a linsleyi Timberlake... . -- ···--- --·---· California Linsley & MacSwain, 1942 > t"' Ashmeadiella SP- -- -- ··--·-··----···-···· --·-· -···-·· -- __ _California new r ecord 0 Callanthiditun illustTe (Cresson) ··-·--·------··California new r ecord ~ Hoplitis pToducta subsp ...... ---·· ····-····-·· --·· California Davidson, 1907 tJj H Hoplitis sambuci Titus _____ ······-··----··· --·· ·-···------California new r ecord 0 Hop litis uvulalis (Cockerell) ·----·········-··--··-·· California new record t"' 0 Osmia laeta Sandhouse ...... California new r ecord 0 Osmia lignaT"ia Say·······-- -·- -·-- ··- --- ···-···· -·--··------California new. record .>< Osmia pikei CockerelL ______California new record < Osmia SP- --·-···------··········-·······-··-············- -· ··-·-- ·- -California new record s: Xylocopa ( ? o?·pifex Smith, not Cresson) .. California Davidson, 1907 ..... _o ...... ,CD t-:> BIOLOGY OF NEMOGNATHA-L INSLEY "\~ D MAc SWAIN 93 upon t he stored pollen, and comp lete their development upon t he contents of a single bee ce ll . In species wi th a wide host range, host differences in the qua nti t ~ · of po ll en stor·ed r esult in con­ siderable size variation in t he adults of N emogncttJw. The fourt h instar is the last feedino· stage and the fi fth instar remains in active wit hin the unruptured skin of t he fourt h instar . 'l'he six t h instar is likewise inactive and remains wit hin the larval skins of the two preceding instars (plate 2, upper fi gures): It has been reported that t he lar val ski n of t he sixth instar is closely adherent to that of the fifth and is not shed at the tim e of pupa­ tion. H oweve r', t hi s i: not always t he ca:e and there is evidence t hat t his feature ma ~ · vary wit hin a species. Lin s l e ~ · and Mac­ Swain (1942 ) have report.ed that in Xemogna.tha sctttell M·is there nu1 ~ · be eit her six or seven lar val instal'S. W hen there are six insta r·s, t he sixth larval skin is shed at the t im e of pupation b11t when the unco mmon seYenth instar occurs it is protected by a to ugh, brown, pupariurn-Jike capsule formed fr·o m t he exuviae of the fourth, fi fth, and sixth la rval instars. In certain ot her species, as .\' . hw·cli lVIacS wain, a ll of t he specim ens reared have had seven insta rs and have pupated in the resistant pupariu m-like capsules. Although most of t he species co mplete their life c ~ ' c l e in a ~ ·ear, t hey a 1·e capable of holding over fOl' two or mo re srnsons.

Nemognatha apicalis J.1eCo nte. This species is active in t he fall and we lun ·e obsened females on man~ · occasions ovipositing on sunflower (FI elia.nthtts a.nm t-ns) (plate 1). Counts of fi fteen ego· masses co llected at random showed a range of eggs per mass from 47 to 285 with a. mea n of 165. This mean is considerably lower than that obsen ed for N. clubia and N. soror (see below). As is true of other species which OYiposit on sunflower·, more than one egg mass may be found on a. single bud, pr es umab l ~ ' as a. result of ov iposition h~' more than one female. 'l'he host of N. apica11:s has not ~ ·et been determined but very likely in volves one or more species of M elissocles which were abundant on sunflo wer·s evel')\vhere we encountered N. apicalis and whi eh commonly C

lake. However, the species which they studied now proves to be N. sctdellat·is LeConte (see below).

Nemognatha cantharidis MacSwain. Oviposition has not yet been observed in this species but the adults frequent flowers of certain desert Compositae, especially Gemea canescens ( 7 collections), and Encelia fat·inosa ( 6 collec­ tions). An interesting feature of its biology was observed by G. A. Marsh who reported to us that he found the species abundant on blossoms of Gemea at night although they were not evident on the same plants in the daytime (locality: 5 mi. S. E. oE Ocotillo, San Diego County, California, March 27, 1949 ) . This nocturnal habit, if it proves characteristic, may well be shared by other desert species of N emognatha and thus explain their relative "rarity" in relation to most other species in the genus. Our largest collec­ tion of N. canthat·idis was made on a partially overcast day near Indio, Riverside County, California.

Nemognatha dichroa LeConte. This species was found ovipositing on Oi1·sium tmdulatum in June and July, 1944, at Vernon, British Columbia, by Hugh B. Leech and on Cirsiunt sp. in Hope Valley, Alpine County, Cali­ fornia, July 18, 1948, by MacSwain. 'l'he infected flowers at Vernon were being visited by M elissodes my sops Cockerell (P. H. Timberlake det. ) which may well prove to be a host of this meloid.

Nemognatha dubia LeConte. An adult of this species was reared from a cell series of M ega chile ( Anthemois) montivaga Cresson, in the dead stem of teasel,

P late 1 Nemognatha CtlJicalis LeConte

Figure 1 (u pper) . Egg masses on back of flower h ead of H elianth1ts anmms (slightly reduced). Figure 2 (lower left). Egg mass ( 9x). Figure 3 (lower right) . Hatching larvae leaving egg mass (9x). BIOLOGY OF NEMOGNATHA-LINSLEY AND MAcSWAIN 95 .96 THE WASMANN JOURNAL OF BIOLOGY, VoL. 10, 1952

( Di1JSacus fullomon) in Putah Canyon, near Davis, California. ~rh e cell series was collected in September, 1940, by Dr. G. E. Bohart. We have found it ovipositing on sunflower (Helianthus annuus) at Romoland, Riverside Cotmty, Califo rnia, July 22, 1946. Five egg masses yielded 115, 293, 357, 375, and 510 eggs _per mass, respectively (mean 330). Davidson (1907) has previously reported rearing thi. · species from cells of Anthiclit~m ema-rgi­ natwn (Say) .

Nemognatha hurdi MacSwain. We have reared this species from mature larvae removed from ·cells of ,lfelissocles ·rob t~sti01· Cockerell in Marsh Creek Canyon, Contra Costa Cotmty, California. The larvae were collected in 1<-,ebruary at which time they were all in the seventh instar en­ dosed in resistant puparium-like exuvial capsules. Adults are active in the fall and we have observed them ovipositing in Oc­ tober on flow ers of Grindelia near Tesla, Alameda County, Cali­ fornia. P. H. Timberlake coll ected an adult on H emizonia lteennannii at Strathmore, Tulare County, Califomia, Septem­ her 30, 1935.

Nemognatha lurida LeConte. Adults of this species were found dead in cells of Megachile occiclentalis 1<-,ox at Loving·, New l\Iexico, in May, 1945, and were observed ovipositing on sunflower (Helianthus annm~s), 5 mi. E . of Acme, Chaves County, New Mexico, August 9, 1950. Although large numbers of egg masses were collected, only two were counted .and these yielded 200 and 375 eggs per mass respectively. Mickel ( 1928) has also reported finding a dead adult of this species in an unbroken cell of. Anthophora occidentctlis Cresson.

Plate 2 N emognatha scutellct1·is LeConte

Figure 4 (above). Sixth instar larvae within exuvial capsule (left) and r emoved from exuvial capsule (right). Figure 5 (lower) . Pupa removed from exuvial capsule (left) aud .adult ( right). BIOLOGY OF NEMOGNATHA- LINSLEY Al'" IJ M.\CSWAIN 97 98 T H E WASMANN JOURNAL OF BIOLOGY, VoL. 10, 1952

Nemognatha lutea pallens LeC onte. Dead, unemerged adults of N. ltdea pallens LeConte, were found in cell series of Anthocopa sp. nesting· in a bank about twenty mil es east of Indio, Riverside County, California, April 1, 1941.

Nemognatha nigripennis LeConte. We have found the typical form of this species ovipositing on E ·l"iogonton fascictda.tum at various localities, occasionall y on Eriophyllttm sp. and other Compositae. P. H . Timberlake has taken adults of the montane dark form on Chr·ysothmnnus sp., Solidago californiw, Erigeron stenophyllu.s, E1·iog01w m elongct­ tum, E. fasciculatum, and E. ttmbellatum, the red desert form on Petalonyx thttrbe1·i, le. ·s commonly on Pluchea. sC?·icea, ()hctenactis ca.?·p hoclinia, and Ba.ileyct m'ltltimdia.tct. Adults of the red form have been reared by us from cells of Jliega chile pmtt1: Cockerell, col lected from a bank nea r Blythe, Riverside County, California, April 2, 1941. Schwarz ( 1903 ) has previously recorded N. nigripennis from a nest of Dianthidium sp. The period of adult activity includes ;\ lay, June, and July.

Nemognatha piezata (Fabricius) . One dead adult specimen of this species was taken from a cell of Anthophom occidentcdi:> Cresson at Loving, Curry County, New iVI exico, in May, 19±5. Later, in the same l oca l it~r , adults were obsened copulating on the nnderside of flowers of Cirsiton and ovipositing on the buds of the sa me plant. Since several larval eo·o· masses had been placed on the underside of almost ever ~r flow er the numbers of primar~ r larvae were unusuall y large. For this reason samples of in c ludin ~ wasps, bees, and butterflies were collected individuall y in an attempt to determine wh ich species carried meloid larvae. Although no larvae were found on either the wasps or butterflies several species of bees had from one to fiv e (average 1.5 ) larvae clinging to their hair::; . Altogether 56 individual bees out of 118 carried larvae principal]~, . beneath the head or on the hairs of the coxae. '!'h e bee s~ec ;

Nemognatha scutellaris LeConte. The habits of this species as a parasite of Jlnthophont linsleyi Timberlake have been briefly discussed elsewhere (Linsley and MacSwain, l 942 ) under the name N. apical is LeConte. Althoug·h N. sctdella1·is is not nearly so widely distributed as some of the other ~pec i es discussed here, it is common along the Pacific Coast where it occurs from sea level to at least !)000 feet elevation. It has a large number of hosts and as a result ex hibit~ a considerable amount of size variation . \Ve have found the adults most fre­ quently on flowers of Achillea rnillefoliurn and various species of Compositae. P. H. 'l'imberlake has taken adults on Erysimtwt asp e n~m, Asclepias sp., Nionarclella exilis, Anisocorna. aamlis, Mala cothTi.r gla.brctta, Eriophyllton lanatwn, and Lc~yia elegans. Their oviposition habits differ from all other species studied in t hat the eggs are inserted in between the florets. Linsley and lHac­ Swain (1942) have noted that in one limited area .V. sctd ellnris and i t~ principal host were visiting different flowers. Th e la rvae primarily infest the cell s of various megachilid bees but have also been recorded b~' Davidson (1907) from Xylocopa, and Linsley and :\facSwain (1942) from AnthOlJhom. . The writers have also found the species under the following conditions: l\Iiner·alking, Tulare County, California. Mature larvae in cell seri es of Osmia pikei Cockerell, Osmic~ sp., and II oplitis uvula/is (Cockerell) all nesting in twigs of elderberry ( Sam b t~cus), Au~-rnst 8, 1!138. These la r-vae were coll ected b~· G. E. Bohart and emerged in the laborator:· as adults in May and June, 1940. \Vood.Y, California. 0Yerwintering, unemerged adults found in cell s of II oplitis sambnci Titus in twigs of eldcrberTy (Sam­ lmcus) . December 28, 1939. :\ft. Diablo, Contra Costa County, California, December 8, 1940. ;\ n overwintering adult enclosed in its exuvial capsule in the 100 THE WASMANN JOUfiNAL OF BIOLOGY, V01 .. 10, 1!!52

terminal cell of Ca1lanthidittrn illustre (Cresson ). An overwin­ tering·, unemerged adult in the subterminal cell of Ashmeadiella sp. in an abandoned nest of Pse1tdomasaris coqnilletti Rohwer. In other cell series of the same species of 1lshrneadiella utilizing aban

Nemognatha soror MacSwain.

Althou~·h this species has not yet been associated witlt its host or hosts, it has frequently bee n obsened on the ftow<'rs of Achillea millej'olinrn. E gg clusters ar·e co mmon]~ · encountered within the flower heads of this plant and a count of six clusters gave a ra11ge from 415 to 7!0 with a mea n of 562 eggs pet· cluster.

Nemognatha sparsa LeConte. 'rhis species was observed at Joseph City, Navajo Count.'', Ari­ zona, August 6, 1950, ovipositing on the unde r·side of buds of G1·indelia.

Nemognatha zonitoides Duges. N ernognatJw zonitoides was coll ected from flowers of (J?-ind elia at Joseph City, Navajo County, Arizona, on August 6, Hl50. It was al.·o found on G1·indelia at Bellview. Curry County, New Mexico on .August 13. Hl:iO.

SELECTED REFERENCES

BALDUF, W. V. 1935. The bionomics of entomophagous Coleoptera. Swift Co., St. Louis, Mo. 220 pp. BIOLOGY OF NEMOGNATHA--L I NSLEY .\XO MA cSWAI N 101

CocKERELL, T. D. A. 1901. Flower and records from New Mexico. Entomologica l News, 12:38- 43. [N emognatha bicolo1· r ecorded visiting fl ower s of Cl eome se1T1t!ata at Las Vegas (p. 40).]

CL . \t ·s ~; N, C. P. 1940. Entomophagous insects. McGraw-Hill. x + 688 pp.

CROS, A. 1912. Entomologie algerienne. N emognatha ch1·ysomelina F. Ses Varietes-Son Evolution. Zeitschrift fiir Wissenschaftliche Insektenbiologie, 8: 137- 141. [Color variation in a dults, flower habits, larva, h ost: Anthidimn discoutale Latreille.]

191 2. S ur )'evolution du N emognatll a c111·ysomeli na F . Bulletin So­ ciete d 'Histoire Naturelle de I'Afr ique du Nord, 3:9-10. [Hosts: Anthidium, M egachile.]

1919. Nemo.Qnatha ch1·ysomelina F . Sa Jarve Primaire. Bulletin So­ ciete d'Histoire Naturelle de l'Afl"ique du Nord, 10 :55- 61. [Egg masses depos ited in flower h eads of E chinops spina­ sus: la rva probably feeds on contents of more than one cell, eating eggs, larvae, a nd honey.]

DAVlDSON , A. c. 1907. from bee cells. Entomological News, 18 : 446. [ N emognatlla dubia recorded from cell of Anthidium emm·ginatum, N . sc1ttellal"is from ( Alcidamw) Hoplitis 711·odu cta and Xylocopa (01·pifex?).]

GRAENlCllEU, S. 1910. On the habits of beetles of the genus Nemognatha. Entomo­ logical News, 21:72-75. [Records a dult Nemognatha v itti ge1·a v isiting Rudbeck i a hi1·ta, less commonly H eZianth1ts occidentalis and Soli­ dago gmrninifolia in Wisconsin ; describes sucking be­ havior on flower s.]

L ECONTE, J. L. 1880. Short studies of North American Coleoptera. Transactions of the America n Entomological Society, 8: 163-218. [Synopsis of Nenwgnatha, pp. 212-215.] 102 THE WASMANN JOURNAL OF BIOLOGY, VoL. 10, 1952

LINSLEY, E. G., and J. W. MACSWAil"'' 1942. The parasites, predators, and inquiline associates of Anthoph­ ora li11sleyi. American Midland Naturalist, 27:402-417, figs. 1-11. [Notes on habits of Nemognatha sctttellaris (as N. ap·i­ calis), pp. 405-406.)

MA CSWAJN, J. "\V. 1951. New North American species of Nemognatha and Zonitis. Pan­ Pacific Entomologist, 27: 72- 80. [ N. h7l1'dii recorded from cells of M elissocles sp.]

MARTIN, C. 1891. [Sur les moeurs des Nemognctthct.] Bulletin Societe Entomo­ logique de France, 1891 : liv. [ N. ch1·ysomelina reared from nest of Anthidi1tnt sp.]

Mrc"EL, Cr.AREN C ~~ E. 1928. The biotic factors in the environmental resistance of Anthoph­ om occidentalis Cress. Entomological News, 39:69- 78. [Dead adult of Nemognatha lurida found in unbroken cell of A_nthophom occidentalis.)

Scnw ARZ, E. A. 1903. [Note on the habits of Nemognatha nig1·ipennis]. P1·oceedings of the Entomological Society of Washington, 5:137-138. [N. nig1'i7Jennis reared from resin nest of ( Anlhitl'iumJ Dianthidittn~ sp. in Arizona.l