vol. 189, no.

2 the american naturalist february 2017

Natural History Note

Living on the Edge: Parasite Prevalence Changes Dramatically

across a Range Edge in an Invasive Gecko

Andrew Coates,1 Louise K. Barnett,2 Conrad Hoskin,2 and Ben L. Phillips1,*

1. School of Biosciences, University of Melbourne, Parkville, Victoria 3010, Australia; 2. College of Marine and Environmental Science,
James Cook University, Townsville, Queensland 4009, Australia
Submitted May 9, 2016; Accepted September 14, 2016; Electronically published December 9, 2016
Dryad data: http://dx.doi.org/10.5061/dryad.gr3p6.

abstract: Species interactions can determine range limits, and

parasitism is the most intimate of such interactions. Intriguingly,
the very conditions on range edges likely change host-parasite dynam-
ics in nontrivial ways. Range edges are often associated with clines in
host density and with environmental transitions, both of which may
affect parasite transmission. On advancing range edges, founder events
and fitness/dispersal costs of parasitism may also cause parasites to be
lost on range edges. Here we examine the prevalence of three species
of parasite across the range edge of an invasive gecko, Hemidactylus fre-
natus, in northeastern Australia. The gecko’s range edge spans the urban-
woodland interface at the edge of urban areas. Across this edge, gecko
abundance shows a steep decline, being lower in the woodland. Two
parasite species (a mite and a pentastome) are coevolved with H. fre-
natus, and these species become less prevalent as the geckos become
less abundant. A third species of parasite (another pentastome) is na-
tive to Australia and has no coevolutionary history with H. frenatus.
This species became more prevalent as the geckos become less abun-
dant. These dramatic shifts in parasitism (occurring over 3.5 km) con-
firm that host-parasite dynamics can vary substantially across the range
edge of this gecko host.
Keywords: Hemidactylus frenatus, invasive, parasite prevalence,
pentastomid, range edge, range expansion.
Introduction
Parasites can profoundly affect population fitness (Blakes-
lee et al. 2012; Dunn et al. 2012; Telfer and Bown 2012), and
where that effect plays out on host range edges, parasites
may cause the expansion or contraction of range edges too
(Case and Taper 2000; Burton et al. 2010; Perkins 2012). Host
range edges are, by definition, clines in host density, and they
often occur where there are clines in the biotic and abiotic en-
* Corresponding author; e-mail: phillipsb@unimelb.edu.au.
ORCIDs: Coates, http://orcid.org/0000-0002-6455-7372; Phillips, http://orcid.org
/0000-0003-2580-2336.
Am. Nat. 2017. Vol. 189, pp. 000–000. q 2016 by The University of Chicago.
0003-0147/2017/18902-56973$15.00. All rights reserved.
DOI: 10.1086/689820
vironment. These concordant clines—in host density, in the
environment, and in the density of intermediate and alternate
hosts—should often lead to profound shifts in host-parasite
dynamics across range edges (Briggs and Hoopes 2004; Cronin
and Reeve 2005). Range edges may also be dynamic—ex-
panding or contracting over time—and we may expect to
see shifts in host-parasite dynamics in these cases too. Even
in the absence of an environmental cline, for example, ex-
panding range edges may lose parasites through serial founder
events or through the lowered fitness and dispersal ability of
parasitized individuals (Phillips et al. 2010; White and Perkins
2012).
Despite the expectation that host-parasite dynamics should
change dramatically across a host’s range edge, there are few
empirical studies demonstrating such patterns (for a rare ex-
ample, however, see Briers 2003). Here we characterize spatial
patterns in parasite prevalence (the proportion of infected in-
dividuals in a population) across the range edge of the Asian
house gecko (Hemidactylus frenatus, a globally widespread in-
vasive) in northeastern Australia. Hemidactylus frenatus was
introduced to Australia in the 1960s but is generally restricted
to urban areas (Hoskin 2011; Vanderduys and Kutt 2013). Re-
cent evidence, however, suggests that populations of this gecko
are beginning to transition into native woodland (Hoskin
2011). This is certainly the case around the city of Townsville
(northeastern Australia), where H. frenatus has been present
since the early 1980s (Barnett et al. 2016). This range edge,
exhibiting a cline in gecko density and spanning a shift from
urban to woodland habitat, is precisely the situation in which
we would expect to observe strong shifts in host-parasite dy-
namics.
The Asian house gecko is host to a suite of parasites, some
of which are new to Australia (Barton 2007, 2015). Indeed,
the possible direct and indirect impacts of exotic parasites
on native species have been a concern regarding H. frenatus
invasion (Barton 2007; Hoskin 2011; Yang et al. 2012). Here
we sample three major macroparasites with very different life
histories (detailed below). We assessed the prevalence of each

The American Naturalist

Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.
000 The American Naturalist
parasite species along transects running across the gecko’s
urban-woodland range edge.
Methods
We sampled Asian house geckos from locations stretching
from inner-urban Townsville (the range core), across the
urban edge (the last buildings in an urban development),
and 2 km into natural woodland. We used seven transects
located around Townsville. Each transect was sampled at
the urban edge and then every 500 m into woodland (out
to 2,000 m). We also assessed an inner-urban transect, with
sites 0, 500, and 1,500 m from the urban edge, although we
could sample only one urban transect due to logistical con-
straints (spatial nonindependence). At each of these 38 sam-
pling sites we estimated the relative abundance of Hemi-
dactylus frenatus, and we collected a sample of geckos (total
n p 231 across the 38 sites), each of which was scored for
the presence/absence of each of three parasite species: Gecko-
bia mites and two pentastomes, Raillietiella and Waddyce-
phalus.
Geckobia bataviensis and Geckobia keegani have both
been found on Hemidactylus around Townsville, although
G. bataviensis is much more common (L. K. Barnett, C. J.
Hoskin, and A. C. G. Heath, unpublished data). Both spe-
cies were introduced to Australia with their host and are
not found on native geckos (L. K. Barnett, C. J. Hoskin,
and A. C. G. Heath, unpublished data). The mites are mostly
parthenogenetic, with males being rare (Rivera et al. 2003).
Geckobia can transmit to hosts via the substrate (Hanley et al.
1995), although transmission via host contact is also a possibil-
ity (Rivera et al. 2003). In either case, we expect Geckobia trans-
mission to be dependent on host density and so decline as host
density declines. We determined the mite infection status of
geckos visually, using a dissecting microscope.
Raillietiella are a genus of pentastomes: hematophagous
crustaceans of the lungs. Raillietiella frenata was likely co-
introduced with H. frenatus (Barton 2007). Adult R. frenata
reproduce sexually in the lungs and shed their eggs with the
host’s feces. Eggs hatch when consumed by a coprophagous
cockroach, which acts as an intermediate host (Lavoipierre
and Lavoipierre 1966; Ali and Riley 1983). If a gecko eats an
infected cockroach, the pentastome larvae migrate to the
lungs and mature. Because of the complex life history, we
predict prevalence to scale with the density of both H. fre-
natus and the intermediate cockroach hosts. All else being
equal, we would expect it to decline as gecko densities de-
cline. We detected R. frenata infection using fecal floats
and searching for eggs using a compound microscope. In
a pilot study, we assessed the effectiveness of our fecal float
technique at detecting Raillietiella infection, using data
from both fecal floats and lung dissections. We had 17 dis-
The American Naturalist
Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.
sected H. frenatus in which we found Raillietiella and also
had fecal samples for analysis. Ten (59%) of these 17 were
infected with mature female Raillietiella, and pentastome
eggs were found in all 10 corresponding fecal floats. This
suggests that our method is close to 100% effective at detecting
female Raillietiella in H. frenatus. In the other seven geckos, we
found only male and/or nymphal pentastomes and, unsur-
prisingly, no eggs in the fecal floats. Thus, if we consider only
the population of adult female Raillietiella, our method
appears both highly specific and highly sensitive. Our data
on Raillietiella should then be considered only as the prev-
alence of adult female Raillietiella.
By contrast to Raillietiella, the pentastome genus Wad-
dycephalus is native to Australia, containing at least five
species (Kelehear et al. 2014). Adults parasitize the lungs
of snakes, while nymphs are found encysted in geckos, in-
cluding (as a novel host) Asian house geckos (Barton 2007).
Waddycephalus is likely transmitted between snakes via
two intermediate hosts: a coprophagous insect and a gecko
(Riley and Self 1981). We detected Waddycephalus infec-
tion in H. frenatus by the presence of nymphs appearing
as characteristic subcutaneous lumps. We predict Waddy-
cephalus prevalence in H. frenatus to be determined by
the abundance of native snakes as well as that of its inter-
mediate hosts: other gecko species and native insects.
Statistical Analyses
Our sites span a sharp transition between urban and wood-
land habitats. Because of dispersal (of hosts and parasites),
however, we would expect variables to respond in a contin-
uous, though stepped, manner along our transects (the step
being due to the sharp habitat transition). To capture this
stepped effect with distance (x), we fit a generalized logistic
function to our data. The function has the following form:
u2l
f (x) p 1 l,
1 1 e(c2x)=s
where the parameters u and l represent the upper and lower
asymptote of a logistic curve, c is the position of the center of
this curve, and s determines the scale of the function. A neg-
ative value of s yields a monotonic decreasing function, a
positive value of s yields an increasing function, and the mag-
nitude of s determines the steepness of the step.
We fitted this function to our data on gecko abundance
and parasite prevalence. Although multiple infections were
common, we took the usual approach of examining parasite
prevalence (i.e., proportion of infected hosts) rather than
intensity (number of parasites per host). For the gecko
abundance data, we assumed a Poisson observation process
(and a log link); for the prevalence data, we assumed a bino-
mial observation process (and so a logit link). The model
 Parasite Prevalence across a Range Edge 000

A Gecko abundance was fitted using the JAGS Gibbs Sampler (Plummer 2013)
25
with minimally informative priors. We initiated three chains
Abundance of H . f renat us
20
and assessed convergence both by eye and by using the
Gelman-Rubin diagnostic. Parameter estimates were taken
from 100,000 samples of each chain following a burn in of
15

20,000 iterations. Convergence was satisfactory in all cases.

All data and analysis scripts have been deposited in the
10

Dryad Digital Repository: http://dx.doi.org/10.5061/dryad

.gr3p6 (Coates et al. 2017).
5
0

Results
−1500 −500 0 500 1500 2000
The relative abundance of Hemidactylus frenatus showed
1.0

B Geckobia prevalence
an unambiguous negative cline with distance outward from
Prevalence of G eck ob i a mites

inner-urban sites (fig. 1A; 95% credible interval for s: 22,809

0.8

to 2330).
Geckobia prevalence was highest at the urban edge (88%)
0.6

and lowest 2,000 m into woodland (43%; table 1). The prev-
alence of Geckobia showed an unambiguous negative cline
0.4

with distance outward from inner-urban sites (fig. 1B; 95%

credible interval for s: 2634 to 23).
Fecal floats detected female Raillietiella frenata with 100%
0.2

accuracy but were unable to detect males. Our prevalence

scores for R. frenata thus refer only to infection by mature
0.0

−1500 −500 0 500 1500 2000

females. Prevalence of these was highest (31%) at inner-urban
sites (table 1). No geckos were found infected with Rail-
1.0

C Raillietiella prevalence lietiella in woodland. The prevalence of Raillietiella showed

Prevalence of female R ai l l i et i el l a

an unambiguous negative cline with distance outward from

inner-urban sites (fig. 1C; 95% credible interval for s: 2721
0.8

to 238).
By contrast to Raillietiella, Waddycephalus prevalence was
0.6

highest (21%) 2,000 m from the urban edge, and we found

no evidence of Waddycephalus in urban geckos (table 1).
0.4

The prevalence of Raillietiella showed an unambiguous pos-

itive cline with distance outward from inner-urban sites
0.2

(fig. 1D; 95% credible interval for s: 4 to 364).

Our model also allowed us to estimate the center posi-
0.0

tion of the clines in prevalence for each parasite species.

−1500 −500 0 500 1500 2000 Interestingly, both of the coevolved parasites (Geckobia
and Raillietiella) show cline centers slightly advanced into
Prevalence of W ad d yceph al us
1.0

D Waddycephalus prevalence
natural habitats (respective 95% credible intervals for Gecko-
bia and Raillietiella: 63 to 793 and 173 to 2,500 m; table 2).
0.8

By contrast, the novel parasite, Waddycephalus, exhibits a

0.6

cline center advanced into urban habitat (95% credible in-

0.4

Figure 1: Hemidactylus frenatus relative abundance (A) and para-

0.2

site prevalence (B–D) at sites spanning the urban edge, from

21,500 m (inner urban) to 2,000 m (natural woodland). In all plots
0.0

we have fitted a generalized logistic curve. In A, the curve is fitted

−1500 −500 0 500 1500 2000 assuming a Poisson observation model (with a log link). The curves
Distance from urban edge (m) in B–D are fitted assuming a binomial observation model (and a logit
link). The size of the circles in B–D indicates host sample size.

The American Naturalist

Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.
000 The American Naturalist

Table 1: Parasite prevalence in Hemidactylus frenatus collected from sites spanning the urban edge, from 21,500 m
(inner urban) to 2,000 m (natural woodland)
Parasite prevalence (%)
Distance from urban edge (m) n Geckobia spp. Raillietiella frenata (female) Waddycephalus spp.
21,500 42 74 31 0
2500 5 80 20 0
0 90 (82) 88 8 6
500 34 (29) 48 0 3
1,000 26 46 0 0
1,500 20 45 0 20
2,000 14 43 0 21
Note: Values of n in parentheses are for Geckobia prevalence.

terval: 22,566 to 2206; table 2). Full details of other param-
eter estimates (upper and lower asymptotes) and model fit
are provided in table 2.
Discussion
Our data reveal distinct spatial patterns in the prevalence
of three parasites of the Asian house gecko, Hemidactylus
frenatus. Both of the gecko’s coevolved parasites (Geckobia
and Raillietiella) appear to have invaded across the urban-
woodland transition but nonetheless show strong negative
clines in prevalence outward from the urban center. By
contrast, the prevalence of the recently acquired parasite,
Waddycephalus, shows a positive cline with distance from
the urban center and some evidence that it is invading into
the urban environment.
The observed decline in Geckobia prevalence is likely
driven by the negative cline in H. frenatus density toward
its range edge: lower gecko densities would be expected to
impair mite transmission. While this local density-dependent
mechanism seems a likely overall driver, our data at distance
zero (where we have high prevalence but low gecko density)
suggest that spatial processes are also important. Here—in

Table 2: Parameter estimates of the logistic function describing the relationship between abundance/prevalence
and distance through transects across the urban/woodland edge
95% credible interval
Variable, parameter Prior Estimate Lower Upper
Gecko abundance (log link):
c N(0, 1,000) 2312.26 21,811.79 1,073.28
l N(0, 1,000) 2.61 23.79 .93
s N(0, 1,000) 21,375.9 22,809.93 2330.35
u N(0, 1,000) 5.02 3.21 8.48
Geckobia prevalence (logit link):
c N(0, 1,000) 336.31 62.66 793.54
l N(0, 1,000) 2.27 21.01 .23
s N(0, 1,000) 2118.09 2634.17 22.66
u N(0, 1,000) 1.63 1.15 2.2
Raillietiella prevalence (logit link):
c N(0, 1,000) 1,186.93 173.71 2,523.04
l N(0, 1,000) 2715.35 22,396.84 210
s N(0, 1,000) 2251.77 2720.83 237.69
u N(0, 1,000) 2.84 21.6 2.06
Waddycephalus prevalence (logit link):
c N(0, 1,000) 21,179.12 22,566.1 2206.52
l N(0, 1,000) 2716.74 21,087.27 2388.17
s N(0, 1,000) 120.24 3.84 363.87
u N(0, 1,000) 22.51 23.17 21.81
Note: The prior was a normal distribution with a mean of 0 and a standard deviation of 1,000 in all cases.

The American Naturalist

Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.

a situation analogous to gene swamping (Kirkpatrick and
Barton 1997; Case and Taper 2000)—we may be seeing a
higher than expected prevalence of parasites because of asym-
metric dispersal of hosts down the density cline, creating a
constant influx of parasites.
The cline in Raillietiella frenata prevalence (and the par-
asite’s absence in woodland sites) may reflect similar den-
sity dependence and spatial processes. It may also, however,
reflect an absence of suitable cockroach intermediate hosts
in native woodland. It is unknown whether native cock-
roaches can carry and transmit R. frenatus larvae. By con-
trast, the American cockroach Periplaneta americana, which
is found in urban Townsville, is believed to fulfil this role
(Lavoipierre and Lavoipierre 1966) and is a species closely
tied to human settlements. Without P. americana, R. frenata
may be unable to obtain a foothold in natural environ-
ments, despite the presence of H. frenatus there (Riley and
Self 1981).
In contrast to the previous two parasite species, Waddy-
cephalus prevalence was significantly higher in woodland
habitat. Native habitat is expected to support a higher
abundance of native snakes (Waddycephalus’s definitive
host) as well as coevolved intermediate hosts. Without its
full complement of hosts, Waddycephalus might be unable
to spread far into urban populations of H. frenatus.
How might these patterns of prevalence influence H.
frenatus invasion? Certainly, parasites impact host fitness;
Raillietiella in particular confer a slew of fitness costs to
lizard hosts (Pence and Selcer 1988; Paré 2008; Caballero
et al. 2015). As the H. frenatus population escapes from
coevolved parasites at its range edge, it is, thus, expected
to benefit from improved fitness. This “enemy release” could
facilitate further spread of H. frenatus into natural environ-
ments as well as potentially exacerbate impacts on native spe-
cies (Keane and Crawley 2002; Blakeslee et al. 2012).
Enemy release could, however, be counteracted by the
acquisition of Waddycephalus. Waddycephalus nymphs mi-
grate through their host’s body and have been observed to
result in direct mortality (G. P. Brown, personal communi-
cation). Thus, Waddycephalus might reduce H. frenatus fit-
ness in native woodland and so have a limiting effect on
range expansion (Levine et al. 2004; Dunn et al. 2012).
Overall, our results confirm expectations of distinct clines
in parasite prevalence across a range edge. The direction of
these clines differs strongly between parasites, likely depend-
ing on parasite life cycle and the density of intermediate and
definitive hosts. These clines in parasitism are driven by fun-
damental aspects of range edges and so may be ubiquitous
well beyond our study system. Our results also point to an un-
derappreciated role that these prevalence clines may have in
regulating range edges, as they may either stabilize a range
edge (via acquisition of novel parasites) or facilitate range ex-
pansion (via escape from coevolved parasites).
The American Naturalist
Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.
Parasite Prevalence across a Range Edge 000
Acknowledgments
We thank D. Barton and G. Brown for their counsel on
pentastome detection and identification. Funding was pro-
vided by the Australian Research Council (DP1094646).
Literature Cited
Ali, J. H., and J. Riley. 1983. Experimental life-cycle studies of
Raillietiella gehyrae Bovien, 1927 and Raillietiella frenatus Ali,
Riley and Self, 1981: pentastomid parasites of geckos utilizing in-
sects as intermediate hosts. Parasitology 86:147–160.
Barnett, L. K., B. L. Phillips, and C. J. Hoskin. 2016. Going feral: time
and propagule pressure determine range expansion of Asian house
geckos into natural environments. Austral Ecology (forthcoming).
doi:10.1111/aec.12416.
Barton, D. 2007. Pentastomid parasites of the introduced Asian
house gecko, Hemidactylus frenatus (Gekkonidae), in Australia.
Comparative Parasitology 74:254–259.
———. 2015. Helminth parasites of the introduced Asian house
gecko (Hemidactylus frenatus) (Gekkonidae), in the Northern Ter-
ritory, Australia. Comparative Parasitology 26:44–55.
Blakeslee, A. M. H., I. Altman, A. W. Miller, J. E. Byers, C. E. Hamer,
and G. M. Ruiz. 2012. Parasites and invasions: a biogeographic ex-
amination of parasites and hosts in native and introduced ranges.
Journal of Biogeography 39:609–622.
Briers, R. A. 2003. Range limits and parasite prevalence in a freshwa-
ter snail. Proceedings of the Royal Society B 270:S178–S180.
Briggs, C. J., and M. F. Hoopes. 2004. Stabilizing effects in spatial
parasitoid-host and predator-prey models: a review. Theoretical
Population Biology 65:299–315.
Burton, O. J., J. M. J. Travis, and B. L. Phillips. 2010. Trade-offs and
the evolution of life-histories during range expansion. Ecology
Letters 13:1210–1220.
Caballero, I. C., A. J. Sakla, J. T. Detwiler, M. L. Gall, T. Behmer, and
C. D. Criscione. 2015. Physiological status drives metabolic rate in
Mediterranean geckos infected with pentastomes. PLoS ONE 10:1–15.
Case, T. J., and M. L. Taper. 2000. Interspecific competition, envi-
ronmental gradients, gene flow, and the coevolution of species’
borders. American Naturalist 155:583–605.
Coates, A., L. K. Barnett, C. Hoskin, and B. L. Phillips. 2017. Data
from: Living on the edge: parasite prevalence changes dramatically
across a range edge in an invasive gecko. American Naturalist,
Dryad Digital Repository, http://dx.doi.org/10.5061/dryad.gr3p6.
Cronin, J. T., and J. D. Reeve. 2005. Host-parasitoid spatial ecology: a
plea for a landscape-level synthesis. Proceedings of the Royal So-
ciety B 272:2225–2235.
Dunn, A. M., M. E. Torchin, M. J. Hatcher, P. M. Kotanen, D. M.
Blumenthal, J. E. Byers, C. A. Coon, et al. 2012. Indirect effects
of parasites in invasions. Functional Ecology 26:1262–1274.
Hanley, K. A., R. N. Fisher, and T. J. Case. 1995. Lower mite infes-
tations in an asexual gecko compared with its sexual ancestors.
Evolution 49:418–426.
Hoskin, C. J. 2011. The invasion and potential impact of the Asian
house gecko (Hemidactylus frenatus) in Australia. Austral Ecology
36:240–251.
Keane, R. M., and M. J. Crawley. 2002. Exotic plant invasions and
the enemy release hypothesis. Trends in Ecology and Evolution
17:164–170.
000 The American Naturalist

Kelehear, C., D. M. Spratt, D. O’Meally, and R. Shine. 2014.
Pentastomids of wild snakes in the Australian tropics. Interna-
tional Journal for Parasitology: Parasites and Wildlife 3:20–31.
Kirkpatrick, M., and N. H. Barton. 1997. Evolution of a species’
range. American Naturalist 150:1–23.
Lavoipierre, M., and M. Lavoipierre. 1966. An arthropod intermedi-
ate host of a pentastomid. Nature 210:845–846.
Levine, J. M., P. B. Adler, and S. G. Yelenik. 2004. A meta-analysis of
biotic resistance to exotic plant invasions. Ecology Letters 7:975–
989.
Paré, J. A. 2008. An overview of pentastomiasis in reptiles and other
vertebrates. Journal of Exotic Pet Medicine 17:285–294.
Pence, D. B., and K. W. Selcer. 1988. Effects of pentastome infection
on reproduction in a southern Texas population of the Mediterra-
nean gecko, Hemidactylus turcicus. Copeia 1988:565–572.
Perkins, T. A. 2012. Evolutionarily labile species interactions and
spatial spread of invasive species. American Naturalist 179:E37–
E54.
Phillips, B. L., C. Kelehear, L. Pizzatto, G. P. Brown, D. Barton, and
R. Shine. 2010. Parasites and pathogens lag behind their host dur-
ing periods of host range-advance. Ecology 91:872–881.
Plummer, M. 2013. rjags: Bayesian graphical models using MCMC.
R package version 3-11.
Riley, J., and J. Self. 1981. A redescription of Waddycephalus tere-
tiusculus (Baird, 1862) Sambon, 1922 and a revision of the taxon-
omy of the genus Waddycephalus (Sambon, 1922), pentastomid
parasites of Asian, Australian and Indonesian snakes, with descrip-
tions of eight new species. Systematic Parasitology 3:243–257.
Rivera, C. M., A. G. Negrón, M. Bertrand, and J. Acosta. 2003.
Hemidactylus mabouia (Sauria: Gekkonidae), host of Geckobia
hemidactyli (Actinedida: Pterygosomatidae), throughout the Ca-
ribbean and South America. Caribbean Journal of Science 39:
321–326.
Telfer, S., and K. Bown. 2012. The effects of invasion on parasite dy-
namics and communities. Functional Ecology 26:1288–1299.
Vanderduys, E., and A. Kutt. 2013. Is the Asian house gecko, Hemi-
dactylus frenatus, really a threat to Australia’s biodiversity? Austra-
lian Journal of Zoology 60:361–367.
White, T. A., and S. E. Perkins. 2012. The ecoimmunology of inva-
sive species. Functional Ecology 26:1313–1323.
Yang, D., E. Gonzalez-Bernal, M. Greenlees, and R. Shine. 2012. In-
teractions between native and invasive gecko lizards in tropical
Australia. Austral Ecology 37:592–599.
Natural History Editor: Mark A. McPeek

“It is generally stated by writers on the Hive-bee that the Oil-beetle (Meloë) is one of its parasites. The possibility that our more common
blister-beetles were similarly parasitic on bees, taken in connection with the frequent complaints from apiarians of the wholesale death of
bees from causes little understood, led me, some years since, to pay attention to the biological characteristics of the blister-beetles, in the
hope of ascertaining whether or not they really bear any connection with bee mortality. From these investigations I am satisfied that Meloë
is only parasitic on the perfect Hive-bee as it is on so many other winged insects that frequent flowers; and that it cannot well, in the nature of
the case, breed in the cells of any social bee whose young are fed by nurses in open cells.” From “On the Transformations and Habits of the
Blister-Beetles (Continued)” by Chas. V. Riley (The American Naturalist 1878, 12:282–290).

The American Naturalist

Downloaded from www.journals.uchicago.edu by University of Cambridge on 01/01/17. For personal use only.