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Life history and population characteristics of the mantid Stagmomantis carolina

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 Life History and Population Characteristics of the Mantid Stagmomantis carolina
(Mantodea: Mantidae)
Author(s): Shonda J. Harris and Matthew D. Moran
Source: Environmental Entomology, 29(1):64-68. 2000.
Published By: Entomological Society of America
DOI: http://dx.doi.org/10.1603/0046-225X-29.1.64
URL: http://www.bioone.org/doi/full/10.1603/0046-225X-29.1.64

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 POPULATION ECOLOGY

Life History and Population Characteristics of the Mantid

Stagmomantis carolina (Mantodea: Mantidae)
SHONDA J. HARRIS AND MATTHEW D. MORAN1
Department of Biology, Hendrix College, 1600 Washington Avenue, Conway, AR 72032

Environ. Entomol. 29(1): 64Ð68 (2000)

ABSTRACT We investigated the life history and populations characteristics of the Carolina
mantid,. Stagmomantis carolina (Johannson), in central Arkansas. Unlike several studies on other
mantid species found in the United States, this species was found at very low densities in the Þeld
as measured by adult abundance and ootheca density. We found that mortality was high during the
egg stage because of developmental failure or egg parasitism. In replicated Þeld plots, mortality and
emigration was high during the 1st and 2nd stadia, resulting in very low numbers of in-plot survivors
(⬍1%) by the 3rd stadium of development. Synchronous versus asynchronous hatching had statis-
tically signiÞcant effects on emigration and in-plot mortality but a nonsigniÞcant effect on in-plot
survival, although the trend was for increased survival in asynchronous plots. Although arthropod
abundance tended to be lower in mantid plots, S. carolina had no signiÞcant effects on total arthropod
abundance, total arthropod biomass, or the abundance and biomass of any arthropod order. Lab-
oratory experiments indicated that this species has slower development during the egg stage and
slower development during the nymph stages compared with related mantid species. This slow
growth may in part explain the high mortality rate and therefore explain the low density of S. carolina
found in this population. However, this growth strategy seems necessary for survival because it allows
for proper timing of oviposition for a univoltine life cycle in central Arkansas. Comparative studies
of this species in other portions of its range would be interesting in determining the generalities of
our Þndings.

KEY WORDS Stagmomantis carolina, praying mantid, community impact, population biology, life
history strategy

THE CAROLINA MANTID, Stagmomantis carolina (Jo-
hannson), is a large generalist predator (50 Ð 65 mm)
found in successional habitats throughout the south-
east United States from the Atlantic coast to the Great
Plains, and from Virginia south to the Gulf Coast.
Unlike several introduced speciesÑTenodera sinensis
Saussure, T. angustipennis Saussure, and Mantis reli-
giosa L.Ñthat have been well studied (Matsura et al.
1984; Hurd and Eisenberg 1984a, 1984b, 1990; Eisen-
berg and Hurd 1990; Fagan and Hurd 1991, 1994; Mo-
ran and Hurd 1996, 1998), little is known about the life
history and ecological interactions of S. carolina.
Throughout much of its range it is univoltine, although
in the extreme southern portion of its range where
there is no killing frost, overlapping generations have
been observed (L. E. Hurd, personal communication).
The development of S. carolina usually involves 6 Ð7
stadia, occurring over an average of 78 d in the labo-
ratory (Rau and Rau 1913, Didlake 1926), but growth
rates in the Þeld are unknown. No data exist on the
mortality rates or community interactions of this spe-
cies.
Before this study, we made Þeld observations on
populations of S. carolina in Faulkner County, central
Arkansas. Hatching appears to occur in April and May
1
To whom correspondence should be addressed.
when 1st-instar mantids are found in the Þeld. We
typically Þnd mantids of various stages of development
during the summer, with adults appearing in the Þeld
by late August. Adults survive until the Þrst killing frost
(usually mid to late November). We, therefore, Þnd 1
generation per year, although occasionally 1st-instar
mantids are observed in late fall (November), pre-
sumably hatching too early and killed soon after by the
Þrst frost. Adult densities are so low that estimates of
populations are difÞcult to determine using mark and
recapture methods. At the end of the season, ootheca
density is also low with a maximum of 0.03 per square
meter measured over 10-ha areas. These densities are
much lower than those observed for other mantid
speciesÑT. sinensis, T. angustipennis, and M. religiosa.
For instance, T. sinensis has a local density up to 2
oothecae per square meter and adult density of 0.6 per
square meter (Eisenberg and Hurd 1990, Hurd et al.
1995).
This study investigated the important mortality
sources during the early development and community
impact of S. carolina in central Arkansas to elucidate
possible reasons for its low density in the Þeld. Vari-
ables studied included egg development rate, nymph
development rate, nymph survival in the Þeld, hatch-
ing pattern and success, and the impact of this pred-
ator on the arthropod community in which it resides.

0046-225X/00/0064Ð0068$02.00/0 䉷 2000 Entomological Society of America

February 2000 HARRIS AND MORAN: LIFE HISTORY OF S. carolina
Materials and Methods
Oothecae Study. S. carolina oothecae were col-
lected from several sites in Faulkner County, AR,
during fall and winter of 1997. These oothecae were
kept in a refrigerator at 2⬚C until March when they
were removed and incubated at 22⬚C. They were
placed in individual 130-ml glass vials, misted occa-
sionally with distilled water, and observed daily for
hatching. Oothecae are often infested by the wasp
Podagrion mantis Ashmead, an egg parasite (Rau and
Rau 1913, Fox 1939). Therefore, we counted number
of mantids that hatched and number of wasps that
emerged. After hatching, we dissected the oothecae to
determine the number of mantids that failed to de-
velop. Using these measurements, we calculated per-
centage mortality for the egg stage of development.
Field Studies. To determine mortality for 1st-instar
mantids and determine the value of asynchronous
versus synchronous emergence, we introduced man-
tids at normal hatching densities to plots in a repli-
cated, controlled Þeld experiment. During March of
1998, 15 plots (2 by 2 m) were established in a 3 ⫻ 5
array. Treatment plots (receiving mantids synchro-
nously or asynchronously) and control plots (no man-
tids) were systematically interspersed within the array
(Hurlbert 1984). Each plot was bounded by a 0.5-m
black plastic barrier stapled to furring strips placed
around the outside edge of the plot. The day of mantid
introduction, a 6-cm band of Tangletrap (Tanglefoot,
Grand Rapids, MI) was painted down the center of the
barrier to trap emigrating mantids and other arthro-
pods. Tangletrap was reapplied as needed to maintain
stickiness. Although many other arthropods are cap-
tured in this type of barrier, the numbers have been
shown to be too low to produce statistically signiÞcant
barrier effects on the arthropod community within the
plots, yet they are effective measures of emigration
(Fagan and Hurd 1994, Moran and Hurd 1994).
First-instar nymphs were introduced into the center
of treatment plots within 24 h of hatching. The mantids
were introduced from 13 April until 11 May. This
rather long period of introduction was because of the
high variation in hatching times for the oothecae. Each
treatment plot received 150 mantids, which corre-
sponds to the emergence number from a large ootheca
(Rau and Rau 1913). The synchronous plots received
all 150 mantids at once, and the asynchronous plots
received 3 introductions of 50 mantids every 5 d. To
make these 2 treatment levels equivalent, we calcu-
lated mantid days by multiplying each mantid by the
number of potential days in the plot. Because the
mantids in the asynchronous plots were introduced in
intervals of 5 d, these plots were maintained for 25 d
whereas the synchronous plots were maintained for
20 d. This time table would equal 3,000 potential man-
tid days for each treatment level (i.e., synchronous
plots: 150 mantids ⫻ 20 d ⫽ 3,000 mantid days; asyn-
chronous plots: [50 mantids ⫻ 25 d] ⫹ [50 mantids ⫻
20 d] ⫹ [50 mantids ⫻ 15 d] ⫽ 3,000 mantid days).
Control plots were maintained for 25 d.
65
Table 1. Egg hatch results from 31 oothecae collected from an
old field
No. % % % fail to Total %
eggs hatching parastized develop mortality
3,666 46.45 3.05 50.49 53.54
During the course of the experiment, we monitored
emigration of mantids by the number captured in the
Tangletrap barriers. This is an effective method for
measuring emigration of mantids (Hurd and Eisen-
berg 1984a). Other species of mantids have exhibited
behavioral effects on spiders (Moran and Hurd 1994,
Moran et al. 1996) so we also monitored the emigra-
tion of spiders out of the plots. Mantids and spiders
caught on the inside half of the barrier were consid-
ered emigrants; any individuals on the outside half of
the barrier were considered to be immigrants and
therefore were ignored. At the conclusion of the ex-
periment, we captured all surviving mantids through
a combination of suction sampling and hand search. To
determine the mantidÕs effect on the arthropod as-
semblage, we used the arthropods captured in the
suction samples. These arthropods were sorted ac-
cording to order and trophic level, counted, dried, and
weighed.
A 2nd Þeld experiment was performed to determine
mortality, emigration, and survival of 2nd-instar man-
tids. There were 4 plots (2 by 2 m) constructed as
described above. Into each plot we introduced nine
2nd-instar mantids that had been raised since hatching
in the laboratory. All mantids were introduced on 17
June to the center of each plot. The plots were
checked daily for emigrants and their instar was re-
corded. The experiment was maintained for 15 d, after
which the plots were suction sampled and hand
searched for surviving mantids.
To estimate growth rates of mantids under optimum
conditions, we raised mantids in the laboratory at
25⬚C. Individual mantids were raised in 130-ml glass
vials equipped with a cotton swab moistened each day
to provide water. Mantids were fed Drosophila mela-
nogaster Meigen ad libitum and monitored daily for
molting.
Statistical Analysis. All of the data sets which vio-
lated the homogeneity of variance assumption were
log-transformed before analysis. Emigration, mortal-
ity, survivorship, arthropod abundance, and spider
emigration in the Þeld experiment were all analyzed
using one-way analysis of variance (ANOVA).
Results
Of the 31 oothecae collected, 8 oothecae failed to
produce any hatchlings. Less than half of the eggs
developed into viable hatchlings; most of the mortality
was caused by failure to develop and a small amount
was attributed to parasitism (Table 1). Of the 23 ooth-
ecae that produced viable offspring, 11 (48%)
emerged asynchronously (i.e., had emergence on ⬎1
d). Average time from the beginning of incubation to
66 ENVIRONMENTAL ENTOMOLOGY Vol. 29, no. 1

the initiation of hatching was 58.3 ⫾ 1.27 (mean ⫾ SE)

d.
By the conclusion of the 1st-stadium Þeld experi-
ment, none of the mantids reached the 2nd instar of
development (duration ⫽ 25 d maximum). However,
in the laboratory, mean time to molt from 1st instar to
2nd was 15.25 ⫾ 0.30 d and no individual required ⬎19
d. The average outdoor temperature during the course
of the experiment was 24.5⬚C, very close to the 25⬚C
temperature maintained in the laboratory. In the 1st-
instar Þeld experiment, the number of mantids emi-
grating during the course of the experiment had
higher total emigration (F ⫽ 17.99; df ⫽ 1, 8; P ⫽ 0.003,
Fig. 1A), and lower mortality (F ⫽ 15.46; df ⫽ 1, 8; P ⫽
0.004; Fig. 1B) when introduced synchronously. How-
ever, the effect of hatching schedule on in-plot sur-
vival was marginally signiÞcant (F ⫽ 4.80; df ⫽ 1, 8; P ⫽
0.060; Fig. 1C) with the trend for higher survival in the
asynchronous plots.
There was a trend for reduced arthropod abun-
dance in both synchronous and asynchronous mantid
plots compared with control plots. However, this
trend was nonsigniÞcant (F ⫽ 1.801; df ⫽ 2, 12; P ⫽
0.207; Table 2). Mantids also showed no signiÞcant
effect on any order of arthropod, trophic level, or
spider emigration (Table 2).
Survivorship for the 2nd-instar experiment was low
(mean, 0.75 ⫾ 0.25, 8%). In contrast to the 1st-instar
experiment, emigration also was low (mean, 0.50 ⫾
0.29, 6%). Therefore, the majority of the mantids died
in the plots (mean, 7.75 ⫾ 0.25, 86%). We found no
individuals that had molted to the 3rd instar by the end
of the experiment, although in the laboratory it re-
quired a mean of 13.89 ⫾ 0.68 d with a maximum of 15 d
to progress from the 2nd to 3rd instar.
The combination of mortality during the egg stage
and 1st 2 instars results in very few survivors. A hy-
pothetical population of 3,000 eggs would result in 1.07
surviving to the 3rd insar according to the mortality
rates for our study.

Discussion
This population of S. carolina exhibits high mortality
in the egg stage, mostly because many eggs fail to
develop and hatch. A minor component of mortality is
parasitism by P. mantis. While this low hatching suc-
cess seems unusual, Rau and Rau (1913) found similar
hatching success and attributed this to infertile ooth-
ecae that may be deposited late in the season. We also
speculate that the long egg developmental time may
cause some eggs to exhaust their energy reserves be-
fore hatching. After hatching, S. carolina nymphs sur-
vive for only 2Ð3 d without feeding (unpublished
data), indicating there are few reserves remaining
after hatching.
We also found that a large number of oothecae
hatched asynchronously and therefore hypothesized
that asynchronous hatching may be a mechanism to
increase survival after hatching. Other mantid species
are known to be food-limited during the early stadia
and exhibit intraspeciÞc competition (Hurd et al.
Fig. 1. Comparison of the mean (⫾1 SE of the mean) of
(A) emigration, (B) in-plot mortality, and (C) in-plot sur-
vival in the 1st-instar Þeld experiment. Emigration, in-plot
mortality, and in-plot survival are measured as total during
course of experiment. *, Statistical signiÞcance (one-way
ANOVA).
1978, Hurd and Eisenberg 1984a), as do many arthro-
pod predators (Rypstra 1983, Wise 1993). This food
limitation results in high starvation rates during the
early stadia. In the Þeld experiment testing mortality
during the 1st stadium, we found that mantids intro-
duced synchronously had higher emigration and
higher in-plot mortality. However, in-plot survival was
marginally signiÞcant and low for both synchronous
and asynchronous mantids (⬍2 per plot average). This
February 2000 HARRIS AND MORAN: LIFE HISTORY OF S. carolina 67

Table 2. Comparison of mean (ⴞ 1SE) arthropod abundance in the 3 treatment levels from the first-instar experiment

Taxa S A C F1,12 P Power

Homoptera 5.80 (1.88) 7.40 (3.19) 14.60 (5.07) 1.35 0.29 0.23
Diptera 2.20 (0.58) 7.80 (1.98) 5.60 (1.63) 3.44 0.07 0.53
Orthoptera 4.80 (2.22) 3.40 (0.75) 18.40 (10.76) 0.30 0.75 0.09
Coleoptera 1.80 (1.79) 0.80 (0.58) 4.00 (6.29) 0.91 0.43 0.17
Araneae 1.00 (0.55) 0.80 (0.583) 1.60 (0.748) 0.43 0.66 0.11
Hymenoptera 0.40 (0.25) 1.40 (0.24) 1.13 (0.31) 1.59 0.24 0.27
Herbivores 15.40 (4.12) 20.40 (4.30) 45.00 (17.11) 2.01 0.18 0.33
Carnivores 2.40 (0.67) 1.60 (0.75) 3.40 (1.29) 0.91 0.43 0.17
Total arthropods 18.00 (4.05) 22.00 (4.95) 48.40 (17.62) 1.80 0.21 0.30
Spider emigration 25.80 (3.29) 27.00 (2.26) 31.40 (3.64) 0.89 0.44 0.17

S, synchronous introduction of mantids; A, asynchronous introduction of mantids; C, control plots.

low survival during the 1st stadium was much lower
than has been found for other mantid speciesÑMantis
religiosa (Fagan and Hurd 1994) and Tenodera sinensis
(Hurd and Eisenberg 1984a). We suggest 2 possible
explanations for this low survival. S. carolina has slower
development (Didlake 1926, this study) than the other
mantid species. This probably exposes S. carolina to
higher predation because small mantids are more vul-
nerable to other arthropod predators (Moran and
Hurd 1994). We also found lower prey abundance at
our Þeld site compared with the studies involving the
other mantid species (Hurd and Eisenberg 1984a, Fa-
gan and Hurd 1994, Moran and Hurd 1994), possibly
causing higher starvation rates.
Survival during the 2nd stadium was low in the Þeld,
as was emigration. This contrasts with at least 1 other
mantid species, T. sinensis, which exhibits high emi-
gration during the 2nd stadium and low emigration
during the 1st stadium (Hurd et al. 1978, Hurd and
Eisenberg 1984a). Mortality remained high, although
somewhat lower, than in the 1st stadium, as would be
expected because larger mantids have fewer preda-
tors, as predation in arthropods is usually size-depen-
dent (Bartley 1983, Hurd 1988).
Stagmomantis carolina seems to face conßicting se-
lective pressures in this portion of its range (central
Arkansas). The growing season extends from March
through October. However, if oothecae are deposited
too early in the season, hatching will occur in the fall,
and the nymphs will be killed during the Þrst frost. We
therefore suggest that this species has a slower devel-
opmental rate to ensure the oothecae are deposited
late enough in the season. S. carolina also exhibits a
long developmental period for eggs; the mean time for
oothecae to hatch was 58.3 ⫾ 1.27 d at 22⬚C. This is
much longer than other mantid species found farther
north (T. sinensis, mean 28 d, unpublished data) where
the growing season is shorter, and may further reduce
the risk of emerging too early. During the course of our
study, we observed 1st-instar mantids in November
and from March until June. Obviously those hatching
in November died soon after. This observation shows
the potential evolutionary pressure on the mantids to
develop slowly, so that they deposit oothecae at the
proper time and hatching occurs in the spring. An-
other option would be the evolution of a bivoltine life
cycle, which we have not observed in Arkansas. S.
carolina would have to reduce its life cycle to ⬇100 Ð
110 d (including ootheca incubation) to acheive this
in central Arkansas, whereas our calculations indicate
the current life cycle of a minimum of 150 d under
ideal conditions (i.e., laboratory).
The result of this strategy of slow development
appears to be high mortality. We believe that mortality
is caused by a combination of predation and starvation
while the nymphs are small. It is known that mortality
probability declines with size in the similar species T.
sinensis, at least in part because of reduced predation
(Hurd and Eisenberg 1984a). We also suspect that S.
carolina suffers high starvation rates because they
were not able to reduce prey numbers signiÞcantly
and prey abundance was low at emergence. Although
the asynchronous hatching we observed may repre-
sent a bet hedging strategy (Partridge and Harvey
1988), it appears to have little positive effect on mantid
populations and may actually increase mortality from
cannibalism between conspeciÞcs (Fagan and Odell
1996). Regardless, the high mortality in the early
stages of development (egg and early stadia) results in
very few mantids reaching adulthood, corresponding
to the low densities of S. carolina we observe through-
out central Arkansas.
This study investigated populations in a small geo-
graphic area (central Arkansas), near the western
edge of the range of S. carolina. This may not represent
optimal habitat for this species, and indeed many spe-
cies exhibit regional differences in population char-
acteristics (Dingle 1981, Brown 1984). For instance,
this portion of the mantidÕs range often has a pro-
longed dry season during the summer, which may
increase mortality from desiccation (Hurd and Eisen-
berg 1984a). A comparative study from other portions
of this mantidÕs range would demonstrate whether our
observed pattern is general or limited to this small
geographic area in Central Arkansas.
Acknowledgments
We thank T. G. Wallace for assistance with these exper-
iments and the Hendrix College Physical Plant for mainte-
nance of the Þeld site. This research was supported by Grant
No. DEBÐ9606880 from the National Science Foundation.
68 ENVIRONMENTAL ENTOMOLOGY
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Received for publication 2 March 1999; accepted 6 August
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