Biology of The Mecoptera

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Ann. Rev. Entomol. 1983. 28:203-28

Copyright 1983 by Annual Reviews Inc. All rights reserved
BIOLOGY OF THE
MECOPTERA
by Royal Melbourne Institute of Technology (RMIT) on 06/27/13. For personal use only.
Annu. Rev. Entomol. 1983.28:203-228. Downloaded from www.annualreviews.org
George W. Byers
Department of Entomology, The University of Kansas, Lawrence,

Kansas 66045
Randy Thornhill
Department of Biology, University of New Mexico, Albuquerque,

New Mexico 87131
One of the minor orders of insects, the Mecoptera comprise about 500
known extant species arranged in 32 genera and 9 families (22, 106). They
have been of interest to entomologists out of proportion to their small
numbers, however, because of their supposed ancestral relationship to the
major orders Diptera and Lepidoptera. Certainly, Mecoptera are among the
oldest neuropteroid or holometabolous insects known from the fossil
record. They make up a relatively conspicuous part of the insect fauna of
the Lower Permian, where the Holometabola first appear (32). The 348
species of Permian, Mesozoic, and Tertiary Mecoptera in fact show more
diversity, based largely on wing venation, than modem ones and have been
assigned to 87 genera in 34 families (171).
Most species of Mecoptera are only rarely encountered by entomologists,
and biological knowledge of them based on observations of the living insects
has been scant until recent years (34). Even morphological studies of any
but the more common species have been few. Purely taxonomic and taxo
nomic-distributional accounts have made up much of the literature on the
order.
Kaltenbach (73) has recently summarized what is known of the Mecopt
era, with emphasis on their morphology, histology, physiology, and behav-
203
0066-4170/83/0101-0203$02.00
204 BYERS & THORNHILL
ior. This is the most complete and thoroughly illustrated publication on the
order. An exhaustive bibliography of the Mecoptera has been compiled by
Schlee & Schlee (131).
GEOGRAPHICAL DISTRIBUTION
Approximate ranges of all mecopteran species were recently indicated

(106). Panorpidae and Boreidae are the dominant families in the northern
continents and Bittacidae in the southern continents. Panorpa includes
about 90 European and temperate Asiatic species and about 50 in eastern

North America and the mountains of Mexico. Neopanorpa, ranging from

India through southern China and Indochina and southward to Java and
Borneo (37), has approximately 90 known species. Leptopanorpa, with 12
species, is limited to Java, although it was formerly thought to be more
widespread (24, 37). Boreus has 10 species in the northern Palearctic region
and 11 in the Nearctic; 2 species of Hesperoboreus and 1 of Caurinus occur
in the western United States (105, 124, 145).
The Bittacidae are most diverse in the Neotropical region, where the
ranges of the small genera Kalobittacus (3 spp.), Pazius (6 spp.), Nannobit
tacus (2 spp.), Neobittacus (2 spp.), Anabittacus (1 sp.), and Issikiella (3
spp.) occur within the broad range of Bittacus. The only widely ranging
bittacid genus is Bittacus, represented in the Neotropics northward from
Paraguay by 16 species, in the Nearctic by 8 species, in the Palearctic and
Oriental regions by 33, and in Africa south of the Sahara by 54.
The single flightless species of Anomalobittacus in southernmost Africa
and of Apterobittacus in central California may have evolved from winged
ancestors within the genus Bittacus somewhere near their present ranges.
In contrast, the recently discovered Californian genus Orobittacus shares a
number of structural peculiarities only with Anabittacus of Chile and Tyt
thobittacus of eastern Australia (166). Twelve species of Harpobittacus and
one each of Austrobittacus, Edriobittacus, and Tytthobittacus are endemic
to Australia. The single species of Hylobittacus is widespread in eastern
North America and apparently has an isolated population in Mexico (30).
Also endemic to Australia are the three genera of Choristidae (2 species
of Chorista, 2 of Neochorista, and 4 of Taeniochorista) (120) and the single
species of Apteropanorpa (family Apteropanorpidae) (35, 119). Nanno
chorista has four species in eastern Australia and two in southern South
America, and its near relative Microchorista inhabits southern New Zea
land. This zoogeographic pattern may help to explain the distribution of
Tytthobittacus, Anabittacus, and Orobittacus mentioned earlier. It is also
the only useful clue in accounting for the oddly disjunct range of the
Meropeidae, with one species of Merope in eastern North America and one
of Austromerope in western Australia (27).
BIOLOGY OF MECOPTERA 205
Brachypanorpa, with two species in the southe rn Appalac hian Moun

tains and two in the mo untainous nort hwe ste rn U ni te d State s, and the fi ve
specie s o f Panorpodes in J apan and Ko re a compri se the family Pano r
podidae. T he remarkab le Notiothauma reedi o f central C hile is the o nly
living me mber of the Eomeropidae (No tiothaumidae) (103, 108, 116, 174).
F amilies, ge ne ra, and species are une ve nly distrib ute d in the vario us
faunal re gions (106). Patte rns of distrib utio n o f e aste rn North Ame ric an
species have been ide ntifie d
distrib utions (25).
Re gio nal taxonomic tre atme nts of Mecopte ra include tho se of No rth
Ame ric a (33), e aste rn North Ame ric a (162, 168), the U SSR (80-82), Japan
(98), C hina (38), Indochina (23), India and adj ace nt re gions (125), I n
done sia (37), Australia (118, 120), and Africa (26, 76).
PHYLOGENY
The phylogeny of Mecopte ra both within the orde r and in re latio n to o the r
holometabolous o rde rs has been the subject of nume rous inve stigations.
Most e arly atte mpts to re late Mecopte ra to Dipte ra, Siphonapte ra, Le pi
dopte ra, and Tr icho pte ra in partic ular were b ase d on a fe w charac te rs, both
primitive and derive d. Hennig (60) showed the importance of using only
unique ly de rived, share d charac te rs (synapo morphie s) as b ase s of taxo
no mic gro ups. Fo llo wing He nnig' s phyloge ne tic me thods, K riste nse n (74)
conc luded that the Dipte ra are the sister group of Mecopte ra + Sipho napt
era. Bo udre aux (14), using the same me tho dology b ut a some what diffe re nt
se t of characte rs, found the Mecoptera to be the siste r gro up of Diptera +
Siphonapte ra. Both these suppose d re latio nships differ from the re sults
re ac he d b y He nnig himse lf (61). T his emphasizes the nee d fo r a phyloge ny
base d upo n as many charac te rs as po ssib le b ut also for the correc t e valu
ation o f these c harac te rs as apomorphous ( derived) o r ple siomorphous
(primitive).
K riste nse n had b ase d his phyloge ny in part on the pre se nce of c utic ular
acanthae in the prove ntriculus of both fle as and ce rtain Mecopte ra, a c har
acte r who se signific ance was stresse d by R ic hards & R ichards (117) and
He pb urn (63) and re ite rate d by Rothschild (121). Bo udreaux (15) showe d
that acanthae actually occur in at le ast te n orde rs of insects, including six
amo ng the Holome tabola. Some o the r similaritie s be tween Mecopte ra and
Siphonapte ra, such as the struc ture of the flage llum of the spermatozoon,
me rit furthe r inve stigation. In mo st insects, the flage llum contains filame nts
(fib rils) so arranged that in cross sec tion they appe ar as two concentric rings
of nine , with two near the center (9 + 9 + 2), where as in the Mecopte ra
and fle as the oute r ring is missing and the formula is 9 + 2 (6, 7, 54).
B acce tti (6) he ld this diffe re nce to be a primitive characte ristic . Departure s
20 6 BYERS & THORNHILL
from the 9 + 9 + 2 patte rn are known in various taxa, and G assne r et al

(54) found that the re are ultrastructural differe nce s in the fl
within the ge nus Panorpa. Multiple se x chromo somes have been found bo th
in the Boreidae and the Sipho napte ra (8, 1 2 1), but in this characte ristic
Boreus differs from o the r inve stigate d Mecopte ra (5).
Ge neral discussions of the po sition of Mecopte ra within the " panorpoid
comple x" of o rde rs c an be found in the works o f Tillyard ( 1 63, 165), Hinton
(68), Kriste nse n (74), Boudre aux (1 5, 1 6), and He nnig (62) .
Suppose d phylogene tic re latio nships amo ng and within the familie s o f
Meco pte ra have been b ase d large ly upon gross mo rpholo gy, both e xtern al
and inte rnal. In rece nt ye ars the wo rks of M ic ko le it (89, 92-94) and W ill
m ann ( 170, 1 72- 174) stand out for their acc uracy o f anatom ical de tail. Both
the se authors, using He nnig' s phylo gene tic me thod (b ut diffe re nt combina
tions o f characte rs), found the siste r gro ups Panorpidae and Pano rpo didae
to be the most de rived or specialize d. Both also e xc lude d the Nannochoris
tidae and Bittac idae from a mono phyle tic group that wo uld include the
othe r se ve n families. They furthe r agree that the Eomeropidae (Notio
thaumidae) m ust be se parate d from the Me ro pe idae and joine d with the
C horistidae, A pteropanorpidae, Panorpidae, and Panorpodidae . In con
trast, Pe nny ( 104) re garded the Me ro peidae as the most ge neralize d family,
followe d b y the Nannochoristidae, C horistidae, and B ittac idae, in that
se que nce. He j udge d Bore idae as c lo se ly re late d to Panorpodidae, large ly
on the b asis of larval similaritie s, with the se two familie s constituting the
siste r group o f Panorpidae . Both M ic ko le it and W illmann have commente d
on He nnig's vie w of phyloge ny o f the Meco pte ra (62) .
We do not inc lude a discussion o f the fossil record he re . Suffice it to say
that the orde r is re lative ly be tte r repre se nted b y fo ssils than are o ther
modem o rde rs. W illmann ( 1 7 1) c atalogue d all the fo ssil species o f Mecopt
e ra, with pertine nt stratigraphic and geo graphic data. M artynova (83) de
scribe d m any ne w Pe rmian forms from the Kuz ne tsk B asin, illustrate d
earlie r de scribe d specie s, and comme nted o n the phyloge ny of the o rde r.
Hennig (6 1, 62) summarize d the record o f Meco pte ra in support of his vie w
o f insect phylogeny.
MORPHOLOGY OF ADULTS
Anatomic al studie s of Mecopte ra have bee n undertake n large ly with a view

to what their results m ight re veal about phylogene tic relationships. The
Pano rpidae (Panorpa) have bee n most thoroughly studie d, b ut re prese nta
tive species in nearly e very fam ily have been e xamine d to some e xtent.
In most adult Mecopte ra, the c lype us and sub genae are conspic uously
prolonge d, forming a rostrum. The m axillary stipite s are corre spondingly
prolonged, lying in an otherwise membranous posterior wall of the rostrum;

the other mouthparts are not particularly elongated (59, 64, 101). In Bra
chypanorpa there is no obvious rostrum (64).
Mandibular structure varies in part according to diet. The mandibles of
phytophagous genera Boreus, Brachypanorpa, and Notiothauma are short,
thick, and have two or more short subapical teeth, whereas those of the
predaceous Bittacidae are slender, flattened, and acutely tipped, with a
single tooth, and cut across each other like scissors [(64, 71, 89, 101);
summarized in (73)]. In the saprophagous Panorpidae, the mandibles are
of an intermediate form (64, 67, 71, 84, 101).

The number of antennomeres varies slightly within a species and among
taxa, e.g. from 16-20 in most Bittacidae and some Boreidae to about 60 in
Chorista and Apteropanorpa. In bittacids the flagellomeres are long and so
slender that their number may be difficult to determine; in other families
they are usually short and thick, ranging from cylindrical (e.g. Panorpa,
Boreus) to nearly moniliform (Merope) (28, 29, 35, 42, 48, 64, 67, 71).
Sensory structures on the antennae of Panorpa and Bittacus have been
described and compared (136).
Internal anatomy of the head of Panorpa communis, notably of the
pharynx and the brain and associated ganglia and nerves, was examined in
detail by Heddergott (59).
Since it clearly displays the sclerites of a generalized neuropteroid ptero
thorax and is large enough to permit relatively easy examination of skeletal
muscles, the thorax of such a mecopteran as Panorpa is a morphology
teacher's delight. There have been a number of good to excellent descriptive
studies of the thoracic exoskeleton, musculature, and wings of Panorpa
(50, 58, 65, 71, 78, 85,86), Bittacus (65, 85, 88, 134, 140), the Boreidae (42,
51, 52, 65, 85), Panorpodidae (71), Meropeidae (65, 69, 85, 87, 172), Notio
thauma (89, 174), and Nannochorista (48).
Conspicuous modifications of thoracic appendages include the reduction
or loss of wings in several families. Wings are absent in Apteropanorpa (35)
and Apterobittacus (73), with both genera showing some reduction of tho
racic size and consolidation or fusion of sclerites. They are reduced to
varying degrees in females of Panorpodidae (28, 36, 71) and are short,
slender vestiges in Anomalobittacus (26). In Boreidae, the wings of females
are short, sclerotized pads, with the forewings usually extending backward
only enough to cover the hind ones (20, 42, 51); the wings of males are
slender, densely sclerotized, hook-like, with spines along the mesal edge,
and are used to grasp the female during mating (40, 42, 51, 124, 141, 175).
Another notable specialization is the raptorial function of the legs of
Bittacidae (134). The structural modifications include a single, large pretar
sal claw (instead of two small ones as in all other Mecoptera), greatly
e lo ngate d tibial spurs, and the ability of the fift h tarsome re to clo se against
the fourt h.
E arly recognition of the taxonomic applic ations o f ge nitalic difference s
be tween species (98,139) focuse d the attention of morphologi sts on ab domi
nal structure as a key to phylogeny (50) and to unde rstanding inte rspecific
re lationships (167). The complic ate d e xte rnal ab dominal anatomy of Notio
thauma has been contrasted w ith that of o the r Mecopte ra and o the r
Holo me tabola (45, 89). Inte rnal abdo minal structure, particularly of the
re pro duc tive syste m, has bee n de scribe d in de tail fo r Panorpa (57,110) and
mo re ge ne rally fo r 9 ther groups (110). Comparative study of the male

ge nitalia re ve als significant structural inconsiste ncie s w ithin the o rde r, such
as abse nce of the spe rm pump (impo rtant in phy logene tic arrangeme nts in
the Mecoptero idea) in Bore idae and Nannocho ristidae (173). M ic ko le it has
recently compare d the female genital anato my in all familie s, with particu
lar atte ntion to the me digy ni um, or "internal ske le ton," poi nting o ut simi
larities and differe nces of phylogene tic rele vance (92, 93). Struc ture and
phylo ge ne tic si gnific ance o f the male no tal o rgan, forme d by parts of ab
dominal terga 3 and 4 to grip the co stal edge of the female's w i ng during
copulation, in Pano rpidae, so me Panorpo didae, and Notiothauma, has been
studie d by Issiki (71) and M ic ko le it (89, 90). Analo gous te rgal mo dific a
tions occur in so me species o f Boreus (95). Ab do mi nal phe romone -di spe rs
ing o rgans have been de sc ribe d in male s of B ittacidae (11,13), Pano rpidae,
and Panorpo didae (147),and their ultrastructure in Harpobittacus has been
e xamine d (46).
Histo logy, lame llar structure, and fracture c haracte ristics of the c uticle
have bee n e xamine d for all nine families (66) although the differe nces found
did not co rre late w i th suppose d phylo ge ne tic re lationships.
EGGS AND EMBRYOLOGICAL DEVELOPMENT
Eggs of Panorpidae, Pano rpo didae, Choristidae, Nannochori stidae, and

Bo re idae are b ro adly ovo id, the e nds about e qually ro unde d (2,3,43,119,
138). At o vipo si tion the chorion i s smooth and o nly lightly scle ro tize d,
usually w hitish to yellow ish b rown i n co lor. In most species of Panorpa, i t
i s cove re d with a fine ne twork o ffi laments produci ng hexagonal o r une venly
poly gonal cells (97, 138). In Panorpa nuptialis, the cho rion lacks this sur
face ne twork and is dark grayish b ro wn (21). B ittacid e ggs such as tho se
of Hylobittacus and Kalobittacus are subsphe rical at ovipo sitio n (30, 99);
in Bittacus and Harpobittacus they are approximate ly c uboidal w he n laid,
w ith the surface s shal low ly impresse d (134). Their c ho rio n i s mo re de nse ly
scle rotize d, may be microsco pically granular, and varies i n color from dar k
yellowi sh brown to dark gray (99, 134). As e mb ryonic deve lopme nt pro-
ceeds, the cuboidal eggs of Bittacus become nearly spherical (134). Egg size
varies between and within species, even among eggs of an individual female
(21, 53). Size increase of eggs during maturation of the embryo has been
noted in Panorpidae, Bittacidae, and Boreidae; this increase may be as
much as 38% in Panorpa (21), 48% in Boreus (141), and 100% in Har
pobittacus (47).
Panorpa and Chorista deposit loose clusters of eggs in preexisting cavities
in moist soil (21,97, 119,164). Boreidae place their eggs singly or in small
clusters in soil around the rhizoids of moss (43, 105) or in epiphytic bryo
phytes (124). Nannochorista lays up to 25 eggs in a row on saturated leaf

litter beneath grasses near water (119). Bittacus drops its eggs more or less
randomly into the ground cover from its hanging position (133, 134).
Duration of the egg stage (oviposition to hatching) varies greatly, depend
ing on the species and to some extent on environmental conditions. In
Panorpa and Neopanorpa, it is 5-10 days for many species (3, 21), but
14--16 days for some species living above 2250 m in Taiwan (177). Panor
podes requires about 26 days (3), and Chorista 12-15 days (19). In contrast,
eggs of Boreus laid in early winter may not hatch until the following spring
(105, 141). A prolonged egg diapause occurs in some species of Bittacus.
Setty (134) reported 216-256 days from oviposition to hatching in Bittacus
stigmaterus under laboratory conditions, but 29-78 days with most larvae
hatching at 32-33 days in nondiapausing B. pilicornis and 20--37 days in
nondiapausing B. punctiger, again under laboratory conditions. Ando (3)
recorded 240-290 days for some diapausing Japanese species.
The only detailed studies of embryonic development are those of Ando
and Suzuki (2-4, 143). Ando traced embryogenesis hour by hour, from first
maturation division to the completion of the blastoderm (2). He found that
eggs of Bittacus have relatively more yolk than those of Panorpa or Panor
podes, which he regarded as a primitive characteristic resembling eggs of
Paurometabola (3). Histology of the embryonic eyes shows them to be
typical compound eyes, quite different in structure from ocelli (4). Rudi
ments of the midgut form adjacent to the stomodaeum and proctodaeum
as soon as these begin to invaginate, then grow together to envelop the yolk
(143). The more general embryological development of Panorpa communis
has been described by Wolf (176).
MORPHOLOGY OF LARVAE
Larval Mecoptera, generally described (e.g. in textbooks) as eruciform,

exhibit a variety of body forms corresponding to the family groups that have
been based on adult characteristics. Those of Panorpidae, Bittacidae, and
Choristidae are in fact eruciform, with a well sclerotized head capSUle,
fleshy but distinctly segmented and single-clawed legs,a sclerotized prono

tal shield, and subconical prolegs on abdominal segments 1-8 (21, 97, 119,
134). Larvae of Boreidae and Panorpodidae, in contrast, are modified
scarabaeiform,with the short,thick abdomen lacking prolegs and usually
curving ventrad,except in Caurinus (43, 105,124). The legs are thick and
fleshy with a slender,tube-like terminal appendage. Those of the prothorax
are directed ventrad,but the middle and hind legs project more to the sides.
Larval Nannochoristidae are somewhat campodeiform,the abdomen being
12-14 times as long as its greatest diameter and having no prolegs (107,
119). The larvae of Meropeidae and Eomeropidae (Notiothaumidae) are

unknown. Those of Apteropanorpidae,though not identified with certainty,

probably are eruciform (49).
Holometabolous larvae as a rule have only simple eyes,or stemmata,but
the lateral eyes of larvae of Panorpa, Neopanorpa, and Chorista are com
posed of 30 or more ommatidia (21,119,177). That these are true com
pound eyes and not clustered simple eyes was determined by Bierbrodt (9)
and confirmed by Ando & Suzuki (4). In the Bittacidae there are only seven
ommatidia,and in the Boreidae usually three (43,105,109),but there are
seven in Caurinus (124). Nannochorista has 16 (68),but the closely related
Microchorista has only a pigmented spot behind the base of each mandible
(107). Brachypanorpa larvae are eyeless (G. Byers, unpublished), as are
those of Panorpodes (N. Suzuki,unpublished).
The thorax and abdomen of panorpid larvae are characterized by small
setiferous sclerites in a generally membranous cuticle. On abdominal terga
8 and 9 there are paired,subconical bases each of which bears a thick but
hollow,annulated,setiform projection; tergum 10 bears a single such struc
ture (21,177). Larvae in the first ins tar possess these enlarged projections
on all abdominal segments. The shapes and arrangement of sclerites and
setae are characteristic of each species. Yie (177) adopted a system of
nomenclature that is used for lepidopterous caterpillars and described and
illustrated first instar larvae of several species of Panorpa and Neopanorpa.
Following this system, Boese (10) was able to construct a key to mature
larvae of 13 North American species of Panorpa.
Bittacid larvae are characterized by a pair of fleshy dorsal projections on
mesonotum, metanotum, and abdominal segments 1-9 (133, 134). Each
projection has three branches,and the shapes of setae on the branches can
be used taxonomically. The projections of abdominal segments 8 and 9 bear
a much thickened seta on the middle branch.
Larvae of Chorista are much like those of Panorpa but lack thickened
dorsal setae except in the first instar (119, 164). In both Boreidae and
Panorpodidae, there is little sclerotization on segments behind the pro
thorax,but sparse setae occur in a characteristic pattern in each genus,and
some species of Hesperoboreus and Brachypanorpa further have transverse

dorsal bands of minute spines on some or all abdominal segments (43; G.
Byers, unpublished).
The elongate aquatic larvae of Nannochoristidae lack dorsal abdominal
processes but have a pair of sclerotized hooks and two slender, membranous
papillae at the end of the terminal abdominal segment (107, 119).
The gross internal anatomy of larvae of Panorpa (56, 109), Bittacus
(134), and Boreus (109) has been investigated. Conspicuous structures are
the large ventriculus about half the body length, the salivary glands near
mid-length of the ventriculus, and the six elongate Malpighian tubules (56).
LARVAL DEVELOPMENT
Larval development in Panorpa proceeds rapidly when food is abundant

and the temperature is favorable. At 20-23C in the laboratory, the fourth
larval stage is attained by Panorpa communis in only 12-16 days, whereas
under natural conditions this may require more than a month (122). Soon
thereafter, the fourth instar larva ceases feeding and becomes quiescent, and
for varying lengths of time the insect remains in this prepupal condition in
a cell that was previously prepared in the soil (21, 122). In the panorpids
and bittacids studied, there is a prepupal diapause; its duration depends
somewhat on the species, but primarily on whether there are one or two
generations per year. In Panorpa species that have both spring and late
summer generations, summer diapause may last somewhat more than a
month and overwinter diapause about 6.5-7 months (G. Byers, unpub
lished). Rottmar (122) found that it was nearly five months in P. communis,
and in Bittacus it has been calculated as about seven months (134). It is not
clear how much of the life cycle of Boreus is spent in diapause; probably
the entire cycle of all boreids requires about two years (43, 141).
Life history observations have been made on Panorpa (21,97, 122, 135,
138, 177), Neopanorpa (177), Bittacus (133, 134), Harpobittacus (47),
Boreus (l OS, 141, 175), Hesperoboreus (43), Brachypanorpa, (36) and Mi
crochorista (107).
PUPAE
Pupation ordinarily occurs in a cell or burrow prepared by the fourth instar

larva in soil or other appropriate substrate. This cell may communicate with
the open air in Bittacus (134), may have an operculum in some Panorpa
species (138), or may have no opening (21). Microchorista sometimes pu
pates in damp moss without making a cell (107), and Caurinus probably
does the same (124). The pupa has much the same form as the adult except
that the wings are tightly folded in their sheaths, the pupal rostrum is not
as long as that of the adult developing within, and the pupal mandibles are
conspicuously larger than their adult counterparts (21, 43, 107, 134, 135,
138, 141, 177). Mecopteran pupae are exarate, can react vigorously to touch
by moving the abdomen and to a slight degree the legs and wings, and can
move their mandibles.
Duration of the pupal stage, from pupal ecdysis to emergence of the
adult, is 10-21 days in Panorpa (21, 122), 15-24 days in Bittacus (133, 134),
14--50 days in Harpobittacus (47), and 37-40 days in Boreus (141). Pilgrim
(107) reared Microchorista from prepupae to adults but apparently did not
time the stadia.
PHYSIOLOGY
Little has been published on the physiology of Mecoptera as such, although

in comparisons of various physiological phenomena in a number of insect
orders Mecoptera have from time to time been represented. Such studies
include physiology of early oocyte-nurse cell development (146), light sen
sitivity of the compound eyes (18), and activity of neurosecretory cells of
the brain (102).
Schwinck (132) examined structural and physiological changes, particu
larly in the cuticle, pericardial cells, and the corpora allata, at various stages
in larval development of Panorpa communis. Some primarily anatomical
studies of larvae of P. communis have touched on physiology in the sense
of general function (56, 109). Functions of various organs in adult mecopt
erans have also been described in largely anatomical treatments. Grell (56),
for example, discussed the development of the greatly enlarged salivary
glands in males of Panorpa and their relation to courtship behavior. Others
have dealt with the physiology of the adult salivary glands, Malpighian
tubules, and fat body in Panorpa, Boreus, and Bittacus (110, 134). Grell
described the anatomy and function of the sperm pump in Panorpa (57),
and Mickoleit (91) discussed sperm transfer by spermatophores in Boreus.
Particularly notable are the numerous studies of aspects of the physiology
of egg development in Panorpa communis by Ramamurty (e.g. 111-115).
Using radioactive tracers and histochemical techniques, he determined
where precursor substances in the haemolymph are converted to carbohy
drates and proteins in the developing egg, where in the egg these are
deposited, and where RNA occurs in the ooplasm.
Not surprisingly, the physiology of winter-active Boreus has been investi
gated (70,137). Unidentified antifreeze substances in the haemolymph per
mit Boreus to supercool and remain active at -6C.
Chromosomes of Panorpa and Bittacus have been studied from sper

matocytes in meiotic prophase. Chromosome numbers vary from n = 20
+ X (0) to n =22 + X (0) in Panorpa and from n = 12 + X (0) to n
= 15 + X (0) in Bittacus [(5) and earlier works cited in (5)]. No chiasmata
form in Panorpa, but homologous chromosomes are joined by a synaptone
mal complex (5, 169). Bittacus has chiasmate meiosis. As indicated, both
Panorpa and Bittacus have XO sex determination. In contrast, Cooper (41)
found that Boreus brumalis has multiple sex chromosomes, males having
an X1X2Y combination and females XIX2'
For more detailed summaries with references on physiology see (55) and
(73).
ECOLOGY AND BEHAVIOR
The ecology and behavior of only a few genera and species of Mecoptera
have been studied in any detail. Many mecopterans are ideal for field studies
of ecology and behavior since the adults are relatively weak flyers, usually
remain near the ground, and often occur in large populations.
Diet and Feeding Behavior

Field studies of Panorpa reveal that soft-bodied dead arthropods (mainly
insects) are the primary food of adults (89-100% of diet) (1 56) . Dead
arthropods are located among the low-growing herbs, where adults feed, by
olfaction (155). Diptera comprise 47-69% of the dead insects in the diet
(156). Panorpa rarely feeds on heavily sclerotized insects, unless the exo
skeleton is ruptured (21, 1 56) . Herb pollen, fruit juices, and perhaps nectar,
and carrion (avian, anuran, and snail) comprise a small component of the
natural diet of Panorpa (98, 156). In the laboratory Panorpa often feeds on
raw meat, fruit, and occasionally on flower petals (21, 53, 73, 79, 156).
Most species of Panorpa are opportunistic in feeding activity, feeding
throughout the day and night (156); however, at least P. nuptialis only feeds
during the day (21). Co-occurring Panorpa species in southeastern Michi
gan overlap almost completely in food type, foraging time, and foraging
location within the herbs; these species rarely forage on the ground or in
the shrub stratum above the herbs (156).
Panorpa exhibits competition for food both within and between species.
This competition is manifested in a complex repertoire of aggressive behav
ior around food, which has been observed by several investigators (21, 53,
153, 156) but has only recently been studied (153, 156). Co-occurring
Panorpa species can be ranked in aggressive dominance on the basis of each
species' ability to exclude the others from food. This ranking is correlated
with size; the largest species win the most aggressive interactions with other
species and the smallest species the fewest (156). Competition for food in
Panorpa influences adult mortality, dynamics of weight loss and gain in
adults, adult dispersal in the habitat, and male sexual behavior.
Competition for food over the evolutionary history of Panorpa popula
tions has resulted in an unusual feeding behavior: feeding to an important
extent on dead insects enswathed in silk in spider webs. This behavior has
been observed in a number of North American species (148, 155; G. Byers,
unpublished) and in European species (100). Panorpa has defenses against
web-building spiders (148) and moves around in webs with little difficulty.
If panorpids become entangled and cannot pull themselves free, often they
regurgitate a brown fluid on the portion of the web entangling them. This
fluid may function to dissolve the web and/or it may may serve to deter
spiders until the insect can free itself. When the regurgitant is smeared on
spiders and other arthropod predators of Panorpa, it sometimes serves as
an effective deterrent. Despite these defenses, web-building spiders are an
important cause of Panorpa mortality. Males of Panorpa feed in webs more
than females. This sex difference is related to mating behavior in that
a significant portion of the food of adult females of Panorpa consists of
nuptial gifts of dead arthropods with accompanying salivary secretions that
are provided by males.
Little is known about the foraging of Panorpa larvae in nature; however,
they are clearly saprophagous, feeding on a variety of dead insects and also
on certain fungi under laboratory conditions (21,53,79). When feeding, the
larvae often occupy a burrow in the soil beneath a source of food (21, 79).
Laboratory-reared adults are atypically small, suggesting an inadequate diet
for normal growth (21).
Adult bittacids are predaceous on a variety of arthropods,mainly insects.
The natural diets of Hylobittacus (99, 149, 150, 156),Harpobittacus (1, 12,
13), and Bittacus (17, 150, 151) have been reported. Studies of Hylobittacus
(156), Harpobittacus (12,13),and Bittacus (151) indicate they capture prey
insects in direct relation to their abundance in the habitat. As in Panorpa,
feeding and mating are usually intimately related. In most bittacids, males
provide arthropods to females as nuptial gifts (13,96, 149). Adult bittacids
can be maintained in a healthy condition on a variety of arthropods under
laboratory conditions (12, 13, 134, 149, 150). Adults of Harpobittacus, in
addition to a diet of arthropods, commonly feed on nectar in nature (13).
The literature on feeding behavior of bittacids contains numerous anec
dotal observations of cannibalism by adults. Cannibalism is largely a labora
tory artifact resulting from crowding and predation on physiologically
weakened individuals and is rarely observed in natural populations of bitta-
cids. In Hylobittacus in nature, conspecific predation infrequently occurs

when the prey is recently emerged and thus soft and vulnerable (154).
Bittacids may grasp prey from a plant or from mid-air while flying. From
a hanging position they use their prehensile tarsi to grasp arthropods that
come into range. They also may sweep the hind legs over the surfaces of
plants, grasping contacted prey (150, 151). Males sometimes steal prey of
other males (150, 151, 153).
Adults of some species of bittacids exhibit nocturnal and crepuscular
feeding activity but others are diurnal (17, 99, 134, 150, 151). Although
bittacids characteristically inhabit the herb stratum, some nocturnally ac

tive species also feed in the tree tops (17, 150, 151).
Bittacid larvae are saprophagous. They have been maintained in the
laboratory on chopped beef, chopped earthworms,or dead insects (46, 133,
134), but also may feed on moss or other plant material (133). Larvae of
Hylobittacus have been observed feeding on dead insects in the field (20).
Both adults and larvae of Boreidae feed on mosses. Lists of moss species
from which boreids have been collected or on which they have been reared
have been published (43, 105, 145). Adults of Brachypanorpa (panor
podidae) are phytophagous,scraping epidermis from soft herbaceous leaves
(28, 36). The larval diet is not known (28).
Little is known about the diets of the remaining five families of extant
Mecoptera. Tillyard (164) supposed that larvae of Choristidae feed on moss,
lichens, and vegetable debris; in the laboratory, adults and larvae fed on
dead insects, fresh fruit, and strained carrots and beef (19; G. Bush, per
sonal communication). The aquatic larvae of Nannochoristidae are preda
tors on larval chironomid midges; the diet of nannochoristid adults is
unknown (63). Adult Notiothauma have been collected by baiting with
rolled oats,suggesting a saprophagous diet
(103); the stomach contents of
Notiothauma are compacted fragments of unidentifiable plants (63). The
diets of larval Notiothauma and both stages of Apteropanorpa and the
meropeids are unknown (63).
Premating and Mating Behavior

Male-emitted pheromones bring the sexes together for courtship in Panorpa
(147), many bittacids (11, 13, 150), and probably Brachypanorpa (147). In
Panorpa the pheromone gland is an eversible pouch in the genital bulb, and
males of Brachypanorpa have a morphologically similar structure. In bitta
cids the pheromone glands are a pair of eversible sacs between abdominal
segments 6-7 and 7-8, although some African bittacids lack the pheromone
structures typical of the family (J . Londt, personal communication). The
pheromones of Panorpa (155), Harpobittacus (11, 13), Hylobittacus (149),
andBittacus (149) are species-specific and operate to attract females from

distances up to 8 m inPanorpa and 15 m in bittacids. The pheromone of
Harpobittacus and Bittacus strigosus attracts both males and females; in
Panorpa. Hylobittacus. and some Bittacus species only females are at
tracted.
Visual signals in the form of wing and body movements are important
in close-range sexual interactions in Panorpa (123, 160, 161) and bittacids
(99, 150, 153, 154). Visual stimuli are apparently important in the sexual
behavior of the Boreidae (43), and the Johnston's organ in the antennae may
detect substrate-borne vibrations (75).

The sexual behavior of Panorpa has been studied in some detail. These
insects are ideal for testing aspects of mating systems theory (153, 160),
sexual conflict theory (161), and life history theory (160). Some general
features of sexual behavior of Panorpa are the following.
There is considerable intrapopulation variation in male mating behavior
in Panorpa. Males exhibit three mating tactics, and individual males are
capable of and often use all three alternatives. Two of these involve nuptial
feeding, i.e. the male presents a food item to the female during courtship
and the female feeds on it throughout copulation. A male also may offer a
female a salivary mass that he secretes. The hard, typically pillar-shaped
mass is attached to a leaf during its secretion. Afterward, the male stands
near the saliva and disperses sex pheromone which attracts females. A
female attracted by the pheromone feeds on the saliva during copulation,
which may last a few hours in some species. Strong sexual dimorphism in
salivary glands, with those of the male comprising a large component of
total body weight, is widespread in Panorpa (56, 110, 160). Alternatively,
a male may locate a dead arthropod, feed on it briefly, disperse sex phero
mone while standing adjacent to it, and then mate with a female as she feeds
on the dead arthropod. During both these kinds of behavior, males display
with wing movements and abdominal vibrations to females attracted to the
pheromone. Males defend nuptial offerings of both types from other males
that attempt to usurp them.
The third behavior used by males is forced copUlation, in which a male
without a nuptial offering rushes toward a female and lashes out his abdo
men at her. A male forcing copulation does not release pheromone but
grasps a leg or wing of the female with his genital claspers. He then re
positions her so as to secure the anterior edge of the female s right forewing
'
in a clamp-like structure on his third and fourth abdominal terga. The male
retains hold of the female's wing with the abdominal clamp throughout
copulation. Although the abdominal clamp of males is used in both forced
and unforced copulations (90, 160, 161), it functions primarily to increase
the pro bability that a mal e c an sec ur e forced copulations when the o th er
two alte rnative s c annot be used (157, 160, 161).
The adoption o f alte rnatives by a mal e Panorpa is dependent o n hi s
ability to compete w ith o th er mal es. Males compete for dead arthropo ds,
which they will feed dir ectly to females and/or feed upon i n order to
produce salivary masses. O nly mal es th at lose in this competi tion exhibit
forced copulation (157, 160, 161).
F emal e fecundity in Panorpa is dependent O n th e alternative used by her
mate. F emales mating with mal es that provide nupti al gifts l ay more eggs
per unit time than females th at experience forced copulation (161), an d

studies offemale choice of mates r eveal that femal es prefer resource-pro vid
i ng male s (153, 160).
Most speci es of Panorpa mate o nl y crepusc ularly and at night (21, 157,
160, 161); however, th ere are species that mate o nl y during the day (21, 73).
Studies of bittacids, espec ially Hylobittacus, h ave l ed to analysis o f some
aspec ts of evolutionary biology and behavio r (149, 151, 153, 154, 158, 159).
For example, the sequenc e of sexual beh avior in Hylobittacus begins when
a mal e acquire s a prey arthropo d. Aft er fee ding briefl y on it, he either
discards it and obtains another i f th e prey is unpal atable or too small o r,
while still holdi ng th e prey in hi s hind l egs, h e initi ates short fli gh ts amo ng
th e her bs. After each fligh t the mal e r el eases pheromone. A responding
female evaluates the prey offer ed by the male by feeding upon it during
cour tship. Female s discriminat e against male s w ith unpalatable or small
prey by flying away and responding to another male's pheromo ne or by
copul ating too briefly for sperm transfer. The mal e t ypic ally retai ns the prey
at th e termination of copUlatio n and, dependi ng o n pr ey size, th e mal e may
use i t in as many as three succ essive copul ations.
Patterns of sexual behavior in Harpobittacus (1, 11, 13) and Bittacus (96,
150, 151) are simil ar to Hylobittacus but may var y i n i mpor tant detail s, e.g.
B. strigosus is the o nly known bittacid to lac k nupti al feeding (17, 150).
Prey theft is an important compo nent o f mal e-male competi tio n for
femal es i n bittac ids (150, lSI, 153, 158, 159). Males have several tactics for
stealing pr ey; in Hylobittacus these i nclude mimicki ng the beh avior o f
females (154).
Some bittacids mate at night a nd others ar e diurnal (96, 150, 151). B.
strigosus mates o nl y at dawn (17, 150).
Borei dae seem to l ac k classic al courtship; they exhibi t unusual sexual
beh avior th at h as been described by a number of workers (40, 43, 73, 95,
105). The greatl y modifi ed w ings of male bor eids are used to po sitio n the
femal e prior to mating and then to hold h er o n th e male's back during actual
mating.
Brachypanorpa (Panorpodidae) mates in the evening, and mating is not

preceded by extensive courtship (36). In the Choristidae males and females
place their mouths together during events surrounding mating (119, 164),
which suggests nuptial feeding via oral secretions. The sexual behavior of
Apteropanorpa, Nannochoristidae, Meropeidae, and Notiothauma is un
known. The elongate male genital claspers of Meropeidae possibly function
in male combat for females. Males of some species of Neopanorpa and
Leptopanorpa (panorpidae) have some elaborate external morphological
features that may also function in male-male competition for females (161).
Oviposition, Sexual Receptivity, and Fecundity

The relationship of mating to female sexual receptivity and egg laying has
been determined in some detail. In Hylobittacus (149) and Bittacus (150;
J. Johnson, unpublished) males introduce a factor into females in the last
few minutes of copulation that turns off female receptivity and turns on
oviposition behavior for durations dependent upon the species. In both
these genera this factor is a product of the male accessory glands. In
Hylobittacus, females after mating become unreceptive for about four hours
during which they lay three eggs. Following mating in Panorpa, females
become sexually unreceptive to males and begin egg laying; the duration of
the nonreceptive period depends on the behavioral pattern of the male with
which the female has mated. Following forced copulation, females of
Panorpa return to sexual receptivity sooner than following copulations in
which males provided resources (161). Males of Panorpa and Bittacidae
apparently are more or less continually sexually receptive.
The oviposition behavior and fecundity of Panorpa and Bittacidae have
been studied in some detail. Females of P. latipennis, observed under labo
ratory conditions, laid different numbers of eggs depending upon the behav
ioral tactics of their respective mates. Females that mated with males who
provided arthropods as nuptial offerings every time the females became
receptive laid about the same number of eggx (x = 3 eggs/day/female) as
females constantly supplied with food (161). There is a strong positive
correlation between female body size and fecundity in Panorpa (161).
Female bittacids typically drop their large eggs singly among the ground
litter while hanging by their forelegs from herbs. In some species, the egg
protrudes from the female's genitalia for several minutes before it is dropped
(99, 134, 158, 159). In Harpobittacus in captivity, the number of eggs laid
by a female varied from 5 to 199 (x = 60 eggs) (13). In Hylobittacus, female
egg output is related to amount of food obtained during mating (149, 158,
159). Setty (133, 134) provided fecundity data for Bittacus in captivity.
The eggs of Boreidae are placed singly or in clumps amid moss rhizoids.
Females lay within a few days (141, 142) to two weeks following first mating
,(43). Hesperoboreus females lay 1.2 eggs per day per female. Svensson (144)
described actual oviposition by a boreid. Oviposition behavior has not been
observed in Meropeidae, Eomeropidae (Notiothaumidae), or Apteropanor
pidae.
Longevity
Adult longevity of Panorpa and certain Bittacidae is related in part to
reproductive biology. This is especially true for males that engage in risky
activities associated with attempts to mate. Males of Panorpa fight vigor

ously and sometimes injure each other, at least under laboratory conditions
(21, 160). Web-building spiders are important predators of Panorpa and
bittacids, and mortality from this predation is significantly male biased.
Male mortality from web-building spiders is size dependent within species
of Panorpa. Small males suffer highest mortality (160) and are least compet
itive for food. They apparently move around more in the habitat and enter
and die in spider webs more often than larger males (160). Males of Hylobit
tacus fly into and die in spider webs more frequently than females owing
to their greater movement associated with obtaining nuptial prey (150, 154,
158). The amount of food females receive from males during mating proba
bly influences female survival because when little or no food is received,
females have to feed on their own and are more exposed to predators.
General competition for food occurs within and between species of adult
Panorpa. This increased movement in the habitat associated with feeding
results in reduced survivorship because of greater exposure to predators,
greater movement into marginal habitats, and considerable weight losses
(156, 160). Mark-recapture studies revealed that longevity of Panorpa
adults is inversely related to the number of competing Panorpa species in
a woods (156, 160). In woods with few species, marked Panorpa lived on
the average 16 days compared to an average of nine days when more
Panorpa species were present. The nature of the interacting species is also
important because Panorpa species differ in their ability to exclude each
other from food. Mortality of the more aggressive species is influenced more
by conspecifics than is mortality of the less aggressive species, which is most
influenced by interspecific competition (156).
Survivorship under laboratory conditions with plentiful food provides
information about potential life span in the absence of normal biological
mortality agents. Vie (177) reported adults of some oriental panorpids
survived for three months. Males and females of two species of Michigan
Panorpa lived up to about 60 days in the laboratory (156; see also 21).
Harpobittacus lived a mean of 15 days in the laboratory (47). Boreus has
been maintained for 51 days (105), and male Hesperoboreus notoperates
lived a mean of 16 days and females a mean of 22 days (43). Given the
polygynous mating systems of mecopterans (160), males are expected to
senesce and die sooner than females, even in the absence of biotic mortality
agents. Males usually die sooner than females in Panorpa (97) and Boreidae
(43) in longevity observations that have been made in the laboratory.
Both sexes of Panorpa vulgaris, under controlled laboratory conditions,
lived about 62 days as adults, but some lived as long as 110 days (39).
Ultrastructural degenerative changes in flight muscle, fat body, Malpighian
tubules, and other organs of the insects appeared after 54 days (39).
Predators and Parasites

The most frequently observed predators of Panorpidae and Bittacidae are
web-building spiders (100, 148, 152, 154, 160). Major aspects of predation
on these Mecoptera by web-building spiders were discussed in the preceding
section. Panorpa and bittacids are also eaten by damselflies, robber flies,
assassin bugs, and roving spiders; Panorpa is also eaten by bittacids (148,
152). Toads (Bufo) and carpenter ants ( Camponotus) eat Panorpa readily
under laboratory conditions and probably are natural predators (R. Thorn
hill, unpublished).
Most records of predators and parasites of Mecoptera are anecdotal.
Boreids have been recovered from the stomachs of trout (43). Other preda
tors of Mecoptera have been mentioned by several authors (see 73). Para
sites of mecopterans include Forcipomyia (Ceratopogonidae), large red
mites, Hymenoptera, fungi, and protozoans [references in (43, 44, 73).
When disturbed, Panorpa adults often fly or drop to the leaf litter where
they remain motionless for a time. Since individuals behaving this way are
difficult to find, this behavior is presumably to avoid predators.
Ecological Distribution
Panorpidae usually frequent moist, northern, continental, temperate forests
with fairly dense, low undergrowth of broad-leaved herbs and shrubs.
Adults occur among the low undergrowth, rarely more than a few feet from
the ground. The larvae and pupae are found in the soil (25, 31, 127, 128,
156, 168). However, there are exceptions to this general pattern. The rufa
group of North American Panorpa species occurs in cultivated fields, pas
tures, and dry, open woods (21). Panorpids penetrate the tropics, but in
such areas they live in cool and forested habitats on mountain slopes (37).
Several species of Panorpa often occur together in the same forest. Stud
ies in Michigan reveal considerable overlap among species in habitat used
by adults (156), without partitioning of the herb stratum by height. Adults
of co-occurring Panorpa species also overlap almost completely in foraging
time and location and general activity pattern. Seasonal differences in emer-
BIOLOGY OF MECOPTERA 22 1
gence time of adults reduce species overlap ( 1 56). Studies of adults of

co-occurring European Panorpa indicate species-specific differences in ver
tical location among the herbs, and the microhabitats of the species differ
in temperature and other physical factors ( 1 28). The European species are
also partially separated by adult emergence time ( 127). Ecological charac
teristics as well as morphological ones were used to differentiate sibling
species ( 129, 130).
Adult and larval Panorpodidae inhabit environments similar to those
typical for montane Panorpa (25, 28, 36). Pupae and mature larvae of
Brachypanorpa have been found in forest soil (28).

Bittacids inhabit forests, grasslands, and occasionally caves (77), but as

in other Mecoptera, high moisture seems to be an important requirement
for most species ( 12, 1 3 , 3 1, 134). Adults typically occur among dense
herbaceous growth, but adults of some species of Bittacus move at night
into the forest canopy where some feeding occurs ( 1 7, 3 1, 150). Bittacid
larvae live in the ground litter, and pupation occurs in the soil (20, 1 3 4).
Adults of Apteropanorpidae and Boreidae are active under conditions of
cold weather. Boreid adult activity is clearly influenced by temperature, the
minimum ambient temperature for activity being close to ODC ( 105, 1 26,
144). Temperature appears to be a more important factor than light in adult
boreid activity (43). Adults and larvae of Apteropanorpidae and Boreidae
live among ground mosses (73), a habitat typically characterized by moist
conditions (124). However, the mosses in which larvae of certain boreids
live may become very dry seasonally. Some European boreid larvae de
scend as much as 20 cm into soil below mosses to reach a humid environ
ment ( 14 1). Larvae of Hesperoboreus form hardened cells in moss, which
may conserve water (43).
The adults of Choristidae and Nannochoristidae occur on broad-leaved
herbs and grasses in moist areas ( 1 1 8, 1 20). The larvae of choristids live
among ground litter ( 1 19, 164), whereas those of Nannochoristidae are
aquatic. Pupation in nannochristids occurs in moist vegetation or soil as in
other Mecoptera ( 107).
Adults of Merope are apparently nocturnal. They have been collected in
hardwood forests and marsh edges, in Malaise, pitfall, and chemical traps,
at lights, and from decaying logs and under rocks (27). The larval habitat
of Meropeidae is unknown. Adults of Notiothauma have been collected in
forests; the larval stage is unknown.
Since most Mecoptera inhibit moist forests, any factor reducing moisture
and/or shade could adversely affect the Mecoptera present. Deforestation
by humans has clearly resulted in extirpation of North American Panorpa
and bittacid populations, and some species are influenced more than others
(25, 1 3 4, 1 56). Human depletion of forests in Java probably has had a
profound effect on the mecopteran fauna there (37). In Mexico, a combina

tion of deforestation and overgrazing appears to have affected present distri
butions of Mecoptera (G. Byers, unpublished).
Abundance
Dense populations of Panorpa (156), Brachypanorpa (27), Hylobittacus
(15S), and Harpobittacus (12) have been recorded. It is unclear whether the
taxa of Mecoptera that are only occasionally represented in museums are
actually rare or simply have not been sought by collectors. Merope tuber,
for example, has only recently been frequently collected, probably because
of widespread use of Malaise traps (27). Other species may be only rarely
observed and collected because of behavioral characteristics such as flying
close to the ground (29) or because of restricted habitats.
Adult Mecoptera often show a distinct peak of population abundance
( 1 56). Males usually emerge before females (43, 1 56), a common pattern in
insects. This, coupled with greater male than female mortality, results in a
predominance of males early in the season (21, 43, 79, 156). Sex ratio of
larvae, pupae, and adult boreids is about 1 : 1 (141). There is no reason to
expect in Mecoptera a sex ratio different from 1 : 1 such as has evolved under
conditions of inbreeding in other insects.
Other Aspects of Ecology and Behavior

In general, mecopterans are relatively weak fliers, and some species have
very reduced wings. Adult dispersal has been studied by mark-recapture
techniques in
Panorpa and Hylobittacus (154, 156); Marked adults of
Panorpa move up to I SO m (x 68 m). Dispersal in these insects can be
=
severely limited by fairly narrow zones of unsuitable habitat. Dispersal in

Panorpa can only be understood in the context of the general ecology of
the insects, because the amount of adult movement depends on the extent
of competition for food and the extent to which suitable habitat is localized
within a woods (127, 128, 156).
Cleaning behavior has been studied in adult Panorpa (72) and Bittacus
(133, 134).
ACKNOWLEDGMENTS
The authors acknowledge with thanks the support of their studies of Me

coptera by the National Science Foundation, currently by grant DEBSO-
22 3 42 to Byers and by grants BNS79-12208, DEB77-01S7S, and
DEB79-10193 to Thornhill.
Literature Cited
1 . Alcock, J. 1979. Selective mate choice cations. Ann. Entomol Soc. Am. 73:
by females of Harpobittacus australis 1 89-96
(Mecoptera: Bittacidae). Psyche 86: 16. Boudreaux, H. B. 1981. About the
2 1 3-17 panorpoid complex. Ann. EntomoL Soc.
2. Ando, H. 1960. Studies on the early em Am. 74:1 55-57
bryonic development of a scorpion fly, 17. Brownson, W. M. 1964. The feeding
Panorpa pryeri MacLachlan (Mecopt and mating behavior of the hangingfiy,
era, Panorpidae). Sci Rep. Tokyo Bittacus strigosus Hagen. MA thesis.
Kyoiku Daigaku 9:227-42 Kent State Univ., Kent, Ohio. 42 pp.
3. Ando, H. 1973. Old oocytes and newly 18. Burkhardt, D., de la Motte, I. 1972.
laid eggs of scorpion-flies and hanging Electrophysiological studies on the eyes
rues (Mecoptera: Panorpidae and Bit
of Diptera, Mecoptera and Hymenopt

tacidae). Sci Rep. Tokyo Kyoiku era. In Information Processing in the Vi
Daigaku 1 5 : 163-87 sual Systems of Arthropods, ed. R.

4. Ando, H., Suzuki, N. 1977. On the em Wehner, pp. 147-53. Berlin!New York:
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Biology of The Mecoptera

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Biology of The Mecoptera

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ANNUAL

Ann. Rev. Entomol. 1983. 28:203-28

Department of Entomology, The University of Kansas, Lawrence,

Department of Biology, University of New Mexico, Albuquerque,

Approximate ranges of all mecopteran species were recently indicated

about 90 European and temperate Asiatic species and about 50 in eastern

North America and the mountains of Mexico. Neopanorpa, ranging from

Brachypanorpa, with two species in the southe rn Appalac hian Moun

from the 9 + 9 + 2 patte rn are known in various taxa, and G assne r et al

Anatomic al studie s of Mecopte ra have bee n undertake n large ly with a view

prolonged, lying in an otherwise membranous posterior wall of the rostrum;

of an intermediate form (64, 67, 71, 84, 101).

mo re ge ne rally fo r 9 ther groups (110). Comparative study of the male

EGGS AND EMBRYOLOGICAL DEVELOPMENT

Eggs of Panorpidae, Pano rpo didae, Choristidae, Nannochori stidae, and

phytes (124). Nannochorista lays up to 25 eggs in a row on saturated leaf

Larval Mecoptera, generally described (e.g. in textbooks) as eruciform,

fleshy but distinctly segmented and single-clawed legs,a sclerotized prono

119). The larvae of Meropeidae and Eomeropidae (Notiothaumidae) are

unknown. Those of Apteropanorpidae,though not identified with certainty,

some species of Hesperoboreus and Brachypanorpa further have transverse

Larval development in Panorpa proceeds rapidly when food is abundant

Pupation ordinarily occurs in a cell or burrow prepared by the fourth instar

time the stadia.

Little has been published on the physiology of Mecoptera as such, although

Chromosomes of Panorpa and Bittacus have been studied from sper

ECOLOGY AND BEHAVIOR

Diet and Feeding Behavior

cids. In Hylobittacus in nature, conspecific predation infrequently occurs

bittacids characteristically inhabit the herb stratum, some nocturnally ac

Premating and Mating Behavior

andBittacus (149) are species-specific and operate to attract females from

detect substrate-borne vibrations (75).

per unit time than females th at experience forced copulation (161), an d

Brachypanorpa (Panorpodidae) mates in the evening, and mating is not

Oviposition, Sexual Receptivity, and Fecundity

activities associated with attempts to mate. Males of Panorpa fight vigor

Predators and Parasites

gence time of adults reduce species overlap ( 1 56). Studies of adults of

Brachypanorpa have been found in forest soil (28).

Bittacids inhabit forests, grasslands, and occasionally caves (77), but as

profound effect on the mecopteran fauna there (37). In Mexico, a combina

Other Aspects of Ecology and Behavior

severely limited by fairly narrow zones of unsuitable habitat. Dispersal in

The authors acknowledge with thanks the support of their studies of Me

of Diptera, Mecoptera and Hymenopt

Daigaku 1 5 : 163-87 sual Systems of Arthropods, ed. R.

Mecoptera from Tasmania. Pap. Proc. 66:21 8-25

cular system of Mecoptera: the thorax. sian)

Univ. Kans. Sci Bull 48:801-44 82. Martynova, O. M. 1959. Mecoptera of

Weibchen (Insecta, Holometabola). II. coptera). Proc. R. Entomol. Soc. Lon

phologie 85:157-64 1 13. Ramamurty, P. S. 1967. The cytoplas

125. Rust M. K., Byera, G. W. 1976. The

Wittenberg: Neue Brehm-Biicherei.

Habitatselektion bei Panorpa commu

graphie der rezenten und fossilen Me

157. Thornhill, R. 1980. Rape in Panorpa von Panorpa communis (Mecoptera).

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