J. Med. Entomol. 1-8 (2015) DOI: 10.1093/jme/tjv127: Ampling Istribution Ispersal
J. Med. Entomol. 1-8 (2015) DOI: 10.1093/jme/tjv127: Ampling Istribution Ispersal
KEY WORDS brown widow spider, Latrodectus, Arachnida, urban entomology, invasive species
Within the spider family Theridiidae, the widow spe- Vetter et al. 2012a) as well as Caribbean islands-Ja-
cies of the genus Latrodectus are well known due to maica (Baerg 1954) and Puerto Rico (R.S.V., unpub-
their notorious bites. Currently, there are 31 accepted lished data). In the Pacific Ocean region, brown
Latrodectus species (Platnick 2014, http://www.wsc. widows are found in locations such as Australia (For-
nmbe.ch/, accessed August 2015) distributed widely ster and Forster 1999), Japan (Ono 1995), Hawaiian
on multiple continents and oceanic islands. The Islands of Oahu, Kaua’i and Hawai’i (Pemberton and
brown widow spider, L. geometricus, C. L. Koch, is a Rosa 1940, Krauss 1944, Bianchi 1945), Marianas Is-
pantropical species that exhibits very low levels of ge- lands of Guam (Kerr 2013), Marshall Islands (Dumb-
netic divergence among individuals even from ex- leton 1957), New Guinea (Levi 1967), the Palmyra
tremely separated localities (Garb et al. 2004). atoll (Krauss 1953), Canton island of the Kiribati
Although initially described from Colombia in South (Degnener and Gillaspy 1955), the Galapagos archi-
America in 1841, the brown widow is most probably pelago (Baert 1994), and the west coast of peninsular
native to the African continent (Garb et al. 2004) and Malaysia (Dzulhelmi et al. 2014) (Fig. 1). Here, we re-
has become established in many tropical and subtropi- port the brown widow’s recent establishment in
cal urban locales. Its distribution includes continental French Polynesia (Tahiti, Moorea, Rurutu), and the
regions as South America (Anderson 1972, Figueira Cook Islands (Rarotonga, Mitiaro, Atiu, Aitutaki).
and Vasconcellos-Neto 1993, Simo et al. 2013), Cen- A life history trait that may aid in the rapid and wide
tral America (Triana et al. 2012), North America dispersion success of the brown widow could be its fe-
(Pearson 1936, Brown et al. 2008, Vincent et al. 2008, cundity; it can produce an egg sac every 3 to 10 d with
an average of 18 to 30 egg sacs in a lifetime (Bouillon
and Lekie 1961, Heeres 1991) and an average between
88 and 149 eggs per egg sac (Baerg 1954, Triana et al.
1
Ingénieur de Rech. en Entomol. Médicale, Laboratoire d’Ento- 2012, Vetter et al. 2012b, Danielsen et al. 2014). The
mologie Médicale, Institut Louis Malardé, BP 30 Papeete- French study of Danielsen et al. (2014) showed a relatively
Polynesia. constant production of eggs, averaging 129 eggs per sac
2
Corresponding author, e-mail: [email protected].
3
Department of Entomology, University of California, Riverside, over the months of June to October from field col-
CA 92521. lected sacs in southern California. If this prodigious
C The Authors 2015. Published by Oxford University Press on behalf of Entomological Society of America.
V
All rights reserved. For Permissions, please email: [email protected]
2 JOURNAL OF MEDICAL ENTOMOLOGY
Fig. 1. Distribution of Latrodectus geometricus and its parasitoid wasp Philolema latrodecti in the Pacific Ocean Region
based on Vetter et al. (2012a), Ono (1995), Simó et al. (2013), Pemberton and Rosa (1940), Krauss (1944), Bianchi 1945,
Pemberton and Rosa (1946), Forster and Forster (1999), Kerr (2013), Dumbleton (1957), Levi (1967), Krauss (1953),
Degnener and Gillaspy (1955), Baert (1994), Bibbs and Buss (2011). The insert map shows the positive areas for Latrodectus
geometricus on the two main islands of the Society Islands (Tahiti and Moorea).
fecundity leads to sufficient survival of spiderlings, some venom components with the thought to be more
L. geometricus could become a significant invasive spe- toxic black widow spiders coupled with its presence un-
cies unless pest management can be implemented in der and around outdoor structures such as flower pots,
newly recorded tropical countries as a means to limit garden facilities and tools commonly used in tropical
population densities. countries, envenomation risk from L. geometricus could
The brown widow spider prefers an urban microhab- be of health concern. L. geometricus is less likely to bite
itat, using man-made objects such as benches, garbage than black widow spiders because it often exhibits a
cans, patio furniture, and flower pots to build webs and death-feigning behavior (thanatosis) when disturbed
retreats in comparison to utilizing the surrounding nat- (McCrone 1964). An African clinical report detailing 15
ural environment, at least in Jamaica and southern Cal- brown widow bites showed that the signs and symptoms
ifornia (Baerg 1954, Vetter et al. 2012a). However, to a were not nearly as severe as that of an African black
lesser extent, it can also be found on vegetation such as widow (L. indistinctus) (Müller 1993). Whereas the lat-
the rosettes of Paepalenthus bromelioides (Figueira and ter had the typical symptoms of latrodectism (e.g., gen-
Vasconcellos-Neto 1993), Eucalyptus trees (Rinaldi eralized muscle and abdominal pain and cramps,
2005), and between web-secured fruit tree leaves (cit- profuse sweating, increased blood pressure, extreme
rus, apple, avocado; Vetter et al. 2012a). Although these restlessness), the two most common brown widow en-
are more natural habitats, they are microhabitats often venomation symptoms were pain on inception of the
associated with human development. Anthropogenic bite and burning pain at the bite site. Nonetheless,
transport equipment like cars, trucks, and maritime brown widow envenomations in some cases can be se-
containers may allow wide distribution of brown wid- vere although generally, bites are mostly benign (Ramia-
ows once they are established (Brown et al. 2008, Simo liharisoa et al. 1994, Cardoso et al. 2003, Kiriakos et al.
et al. 2013). 2008, Goddard et al. 2008, Almeida et al. 2009, Suchard
Latrodectus spiders commonly feed on insects, arach- 2009). Until there is a more complete understanding of
nids, and even small vertebrates like geckos, other liz- brown widow envenomation effects in humans, their
ards, and mice using venom that can cause muscle bites should be treated with concern. In addition to the
injuries and have neurological and proteolytic activities toxic effect, there is also the psychological trauma that is
like those caused by L. geometricus toxins (Reyes-Lugo induced (often self-induced) because the spider involved
et al. 2009). Because it is a widow spider that may share is a member of the infamous widow spider genus.
2015 MARIE AND VETTER.: BROWN WIDOW SPIDER IN FRENCH POLYNESIA 3
Table 1. Latrodectus geometricus specimens collected (first specimen in 2006) and surveys between 2012 and 2014
Tahiti Pai horo 2014 17 430 48.3100 S Waste storage 8 46 0.5–1 m
149 200 7.7700 W
Tahiti Motu Uta Intl wharf 2012–2014 17 320 14.3200 S Waste storage 119 243 1–2 m
149 340 45.3000 W
Tahiti Arue 2013–2014 17 310 41.0000 S Container waste 23 79 0–05 m
149 320 13.3000 W
Tahiti Arue 2012 17 320 1.9700 S Flower pots 5 6 0–05 m
149 320 1.7900 W
Tahiti Pirae 2013 17 310 39.1500 S Flower pots 1 2 0–05 m
149 320 55.8900 W
Tahiti Punaauia 2012–2014 17 370 57.9200 S Waste storage 39 86 1–2 m
149 350 27.5800 W
Tahiti Faaa 2013 17 340 14.4300 S Waste storage 2 4 1–2 m
149 370 3.8100 W
Tahiti Faa’a St Hilaire 2013 17 340 17.2500 S Under car 9 15 0–05 m
149 350 59.8800 W
Tahiti Mahina 2012 17 300 12.6600 S Furniture 1 3 0.5–1 m
149 290 22.4400 W
Tahiti Pirae 2013 17 320 47.0500 S Christmas tree 1 0 0–05 m
149 320 51.9100 W
Moorea Pihaena 2006 17 290 15.5700 S In house 1 0 0.5–1 m
149 490 43.3600 W
Moorea Temae 2014 17 280 53.4400 S Waste storage 10 22 0–05 m
149 460 34.0500 W
Rurutu Teanapapa 2013 22 270 5.5800 S Table bunch 4 5 0.5–1 m
151 220 34.2200 W
Rarotonga Airport and Wharf 2014 21 120 14.7500 S Containers NR 66 NR
Cook Island 159 470 1.6300 W
a maritime shipping container (6 by 2.4 by 2.6 m3) com- Table 2. Microhabitats of Latrodectus geometricus in French
ing from the international wharf of Los Angeles. Soon Polynesia (223 female spiders in 13 positive locations)
afterward, a resident of Tahiti captured a female brown
widow in an artificial Christmas tree; this spider was Category Location Percentage %
misidentified as a black widow on social media because
Car / transportation Under car, trucks . . . 9.87
of its shape, dark coloration and ventral orangish-red garden items Furniture and artificial plant 0.90
hourglass. Additional specimens that have been col- Flower pots 2.69
lected on the main island of Tahiti during the year after Bunch/ table/ chair 2.24
discovery were identified. Our surveys since this time Waste treatment Waste container 65.47
company Under paper waste storage grid 11.21
revealed the establishment and distribution of the brown Vertical wall 4.04
widow on Tahiti, Moorea and Rurutu islands of French Under sewer grid drain cover 3.59
Polynesia. We were informed of additional establish-
ment of the spider in the Cook Islands: Rarotonga in
June 2014 (G. MacCormack, personal communication),
Mitiaro and Atiu in December 2014, and Aitutaki in A total of 511 egg sacs from 223 females in nine
February 2015 (M. Poeschko, personal communication). areas were dissected to estimate the egg clutch size of
Regarding brown widows, we collected 223 females, L. geometricus in French Polynesia. From the 511 egg
35 males and 511 egg sacs from 13 of the 30 sites in sacs, only 129 (25.2%) contained viable eggs and/or
French Polynesia that were inspected (Table 1, Fig. 2). well-developed spiderlings, 203 (39.7%) were empty
Of these, 84.3% of the spiders were found within areas with successful emergence, 21 (4.1%) were not viable
related to waste recycling centers with 65.5% of the fe- (eggs sterile or coated with fungus), and 158 (30.9%)
male spiders located inside, underneath or on top of contained parasitoid wasps.
portable waste containers. Only 9.9% were located un- The Kruskal–Wallis test showed that egg sac clutch
derneath automobiles and 5.8% were found in associa- size (eggs plus spiderlings) was significantly different
tion with garden and picnic items (Table 2). Brown for at least one of the locations (v2 ¼ 32.59, df ¼ 4,
widows were absent in natural and rural places, prefer- P < 0.0001, Table 3). Multiple comparison techniques
ring urban areas instead. In regard to height, 75% of determined that the egg clutch size for the Cook Is-
the specimens were collected between 1 to 2 m from lands was significantly smaller than that of French
the ground in areas rarely frequented by humans Polynesia (field and lab) and California 2011 and 2012
(waste industries). On the other hand, the remaining (Table 3). In addition, French Polynesia (lab and field)
25% that were found under 1 m height were from the was significantly smaller than that from California
underside of automobiles, flower pots, plastic table or 2012. French Polynesia (field) was determined to be
chairs outside the houses. significantly different from California 2011.
2015 MARIE AND VETTER.: BROWN WIDOW SPIDER IN FRENCH POLYNESIA 5
Table 3. Means and standard deviations of the egg clutch size for 1) the entire sample at five locations or years and 2) a randomly cho-
sen subsample of 21 egg sacs from the four largest samples for statistical comparison of equal sample sizes
All Subsample of 21
Fig. 3. Positive Latrodectus geometricus stations and Philolema parasitism on egg sacs for Tahiti (Mahina to Paihoro),
Moorea Islands, Rurutu Island, and Cook Islands. Numbers at the top of the histobars are the sample sizes.
Egg sac hatch success for French Polynesia (number contained 4,015 wasps (v ¼ 25 wasps per egg sac) with
of empty egg sacs þ viable egg sac/ all egg sacs) was 98.1% of the egg sacs being completely parasitized.
limited to 65% due to parasitism. The parasitoid wasps
found in 158 egg sacs were identified as Philolema
Discussion
latrodecti. Parasitoid wasp infestation was found in
brown widow egg sacs in five of 11 locations on Tahiti The brown widow spider has colonized many areas
(Fig. 3) but none were found on Moorea or Rurutu of in the Pacific Islands and is becoming widely distrib-
French Polynesia or any of the Cook Islands. Regard- uted in two new areas in the South Pacific. The brown
ing the Tahitian egg sacs, 31% had parasitoids including widow spider has now established itself on three islands
those from four closely situated cities (Arue, Papeete, of French Polynesia and four of the Cook Islands, add-
Faa’a, Punaauia) and one isolated station of Paihoro, ing a new theridiid species to the last arachnid inven-
which is a landfill site in a rural habitat. Indeed, be- tory for French Polynesia (Dierkens and Charlat 2011).
cause trucks from Papeete empty their containers in This dispersion in a short period suggests that shipping
Paihoro, some spiders can get knocked loose from the containers are the most probable pathway for the
containers when they are emptied and colonize these brown widow spider to invade new territories.
new areas. In French Polynesia and the Cook Islands, the sam-
When emerged P. latrodecti wasps of 3 d of age pled areas with brown widows are mainly associated
were introduced to 2-d-old egg sacs, 100% of the spi- with recycling centers. Therefore, this would indicate a
der eggs were parasitized in laboratory conditions (in- low-level envenomation risk due to the limited distribu-
side a Petri dish with six egg sacs), producing 312 tion of the brown widow, restricted mostly to recycle
wasps (147 females and 165 males, average: 52 wasps center containers of inland and international maritime
per egg sac). The time of development from eggs to traffic. However, eventually new shipping containers
adult emergence took roughly 16 d (15.8 6 1.0) at 27 C departing from Tahiti may be contaminated with egg
and 70% RH in the environmental test chamber. The sacs and spiderlings, further spreading this invasive
158 parasitized field-collected egg sacs from Tahiti species to new areas. Anderson (1972) describes how
6 JOURNAL OF MEDICAL ENTOMOLOGY
the brown widow and plants in which it prefers to nest some of the brown widows that originally infested
had thoroughly recolonized, in 3 mo time, a section of French Polynesia, it is unlikely that this is also the
Brazilian beach that had been burned. Private resi- source of the Philolema wasp, as Vetter et al. (2012b)
dences in French Polynesia may be infested from pub- dissected over 4,000 Latrodectus egg sacs from many
lic service vehicles, which may inadvertently spread the locations in southern California and found no Philolema
spider around the islands. Homeowners need to exert wasps.
caution when moving objects outside their homes or Risk of human envenomation by brown widow spi-
when making repairs underneath cars. Because of the ders is low for French Polynesia because, for the mo-
pantropical distribution of L. geometricus, its origin in ment, brown widow populations are most plentiful in
French Polynesia and the Cook Islands could have oc- areas with restricted access by the general public.
curred from a number of potential maritime exporting Nonetheless, the brown widow should not be ignored
areas such as Australia, Japan, Hawaii, or California. considering its preference for urban areas especially
The egg clutch size of brown widows in this study around homes (Baerg 1954, Vetter et al. 2012a). Edu-
was low compared to other studies around the world. cation about the spider should not be suppressed, as it
From this study, egg clutch size was significantly could be an envenomation threat during dengue out-
smaller for subsamples of egg sacs from either French breaks and vector control campaigns when exposure to
Polynesia or the Cook Islands in comparison to those brown widows may increase as people handle flower-
from southern California in 2011 and 2012 (Table 3). pots, removing water in attempt to reduce mosquito-
Egg sac clutch size from Costa Rica (141.9 6 62.4, no breeding sites.
max given, N ¼ 47; Triana et al. 2012) was in the range Although mosquito-vectored illnesses pose a greater
of the high California data although that publication threat to the populace of French Polynesia than spi-
does not list individual raw data for statistical compari- ders, because there are no other medically important
sons. Only in Jamaica (mean ¼ 88, N ¼ 11; Baerg 1954) spiders on the islands, people may overreact to the
did egg clutch numbers resemble those produced by news of a possible toxic spider in the area. Overreaction
our data here. In addition, Baerg mentions that the Ja- is typical in response to widow spiders no matter how
maican brown widow egg sacs were collected when the minor the threat or how much reassurance is provided
rainy season was well underway. Perhaps for Tahiti as by public education (R.S.V., personal observation).
well as other tropical regions, the number of eggs per
sac or the number of sacs per female varies with wet or
Acknowledgments
dry season, which would be influenced by food avail-
ability. However, although rain is usually thought to be We are grateful to the owners of properties sampled on Ta-
beneficial to insects because of increased productivity hiti for providing live specimens at the start of the study, es-
of the flora on which they feed, rain could also be det- pecially E. Claridge who discovered the first specimen in
rimental by restricting feeding and growth of insects Moorea (2006) and sent samples from Rurutu (2013). M. A.
during extended wet periods. Cheong Sang assisted on the field missions. We thank our col-
leagues G. MacCormack and M. Poeschko who reported and
The Tahiti brown widow population appears less
sent egg sac specimens from Rarotonga (Cook Islands) for
invasive than that of the Cook Islands or southern Cali- this study, R. Putoa who alerted us by providing the 2012
fornia (Vetter et al. 2012a). The lower Tahitian brown specimen from a shipping container (biosecurity), and the
widow populations can possibly be attributed to the ex- Service of Development Rural (SDR), which provided loca-
clusive presence of parasitoids on Tahiti in comparison tions for the first specimens from private residences and
with surrounding islands. home owners. Linda Penas (UC Riverside) provided statisti-
Parasitized egg sacs contribute to population limita- cal analysis. Finally, we thank the local television and press
tion, but it is unclear why parasitoid wasps are media releases, which alerted the general public to the pres-
abundant in some locations compared with others. ence of the spider, which in turn resulted in a greater number
However, even if brown widow populations seem to be of spider submissions from different locales.
detrimentally affected by a parasitoid, some locations
did not show any reduction of populations from these References Cited
controlling influences, such as under cars or in flower
Almeida, R.A.M.B., R. S. Ferreira Jr., C. R. Chaves, and B.
pots. In French Polynesia, P. latrodecti may have a sea- Barraviera. 2009. Envenomation caused by Latrodectus
sonal activity peak and habitat preferences that should geometricus in Sao Paulo State, Brazil: A case report.
be studied to understand whether it can control and J. Venom Anim. Toxins incl. Trop. Dis. 15: 562–571.
limit the distribution of L. geometricus. Anderson, M. P. 1972. Notes on the brown widow spider,
The distribution of P. latrodecti is poorly known but Latrodectus geometricus (Araneae: Theridiidae) in Brazil.
appears to be disjunct and spread over a wide area. It Great Lakes Entomol. 5: 115–118.
is known from Florida (Bibbs and Buss 2011), Hawaii Baerg, W. J. 1954. The brown widow and the black widow spi-
(Fullaway 1953), the Marshall Islands (Schreiner 1989), ders in Jamaica (Araneae, Theridiidae). Ann. Entomol. Soc.
Australia (Lotfalizadeh et al. 2007) and Micronesia Am. 47: 52–60.
Baert, L. 1994. Notes on the status of terrestrial arthropods in
(Herting 1971). A Philolema species was noted as being Galápagos. Noticias de Galapagos 54: 15–21.
collected from brown widow eggs sacs in Costa Rica Bianchi, F. A. 1945. Notes on the abundance of the spiders
but no specific epithet was provided (Triana et al. Latrodectus mactans, Latrodectus geometricus and Argiope
2012). Although maritime shipping containers from the avara, and their parasites on the Island of Hawaii. Proc. Ha-
port of Los Angeles, CA, could have been the source of waiian Ent. Soc. 12: 245–247.
2015 MARIE AND VETTER.: BROWN WIDOW SPIDER IN FRENCH POLYNESIA 7
Bibbs, C. S., and L. J. Buss. 2011. Widow spider parasitoids, Krauss, N.L.H. 1944. Notes on insects and other arthropods
Philolema latrodecti (Fullaway) (Insecta: Hymenoptera: from the islands of Molokai and Maui, Hawaii. Proc. Hawai-
Eurytomidae) and Baeus latrodecti Dozier (Insecta: Hyme- ian Entomol. Soc. 12: 81–94.
noptera: Platygastridae). University of Florida, Publication Krauss, N.L.H. 1953. Insects and other invertebrates from Pal-
#EENY 515. myra Atoll and Christmas Island. Proc. Hawaiian Entomol.
Bibbs, C. S., A. M. Vitoreli, G. Benny, C. L. Harmon, and R. Soc. 15: 217–220.
W. Baldwin. 2013. Susceptibility of Latrodectus geometricus Levi, H. W. 1967. Cosmopolitan and pantropical species of
(Araneae: Theridiidae) to a Mucor strain discovered in north theridiid spiders (Araneae: Theridiidae). Pac. Insects 9:
central Florida, USA. Fla. Entomol. 96: 1052–1061. 175–186.
Bouillon, A., and R. Lekie. 1961. Cycle and rhythm in the ovu- Levi, H. W., and D. E. Randolph. 1975. A key and checklist of
lation of the spider Latrodectus geometricus C. Koch. Nature American spiders of the family Theridiidae north of Mexico
191: 620–621. (Araneae). J. Arachnol. 3: 31–51.
Brown, K. S., J. S. Necaise, and J. Goddard. 2008. Additions Lotfalizadeh, H., G. Delvare, and J. Y. Rasplus. 2007. Phylo-
to the known U.S. distribution of Latrodectus geometricus genetic analysis of Eurytominae (Chalcidoidea: Eurytomidae)
(Araneae: Theridiidae). J. Med. Entomol. 45: 959–962. based on morphological characters. Zool. J. Linn. Soc. 151:
Cardoso, J.L.C., A. D. Brescovit, and V. Haddad, Jr. 2003. 441–510.
Clinical aspects of human envenoming caused by Latrodectus Lotz, L. N. 1994. Revision of the genus Latrodectus (Araneae:
geometricus (Theridiidae). J. Venom Anim. Toxins incl. Trop. Theridiidae) in Africa. Navors. Nasionale Mus. Bloemfontein
Dis. 9: 418. 10: 1–60.
Danielsen, D.W.R., D. E. Clarke, S. J. Valle, A. A. Anselmo, McCrone, J. D. 1964. Comparative lethality of several Latro-
L. S. Vincent, and R. S. Vetter. 2014. Natural egg sac dectus species. Toxicon 2: 201–203.
clutch size of the brown widow spider, Latrodectus geometri- Müller, G. J. 1993. Black and brown widow spider bites in
cus (Araneae: Theridiidae) in southern California. Bull. So. South Africa: a series of 45 cases. S. Afr. Med. J. 83: 399–405.
Calif. Acad. Sci. 113: 100–102. Ono, H. 1995. Records of Latrodectus geometricus (Araneae:
Degnener, O., and E. Gillaspy. 1955. Canton Island, South Theridiidae) from Japan. Acta Arachnol. 44: 167–170.
Pacific. The Pacific Science Board, Atoll Res. Bull. 41: 55. Pearson, J.F.W. 1936. Latrodectus geometricus Koch, in south-
Dierkens, M., and S. Charlat. 2011. Contribution à la connais- ern Florida. Science 83: 522–523.
sance des araignées des ı̂les de la Société (Polynésie Pemberton, C. E., and J. S. Rosa. 1940. Notes on the life his-
française). Revue Arachnol. 17: 63–81. tory of Baeus californicus Pierce, an egg parasite of the black
Dumbleton, L. J. 1957. Parasites and predators introduced into widow spider. Hawaii. Planters’ Rec. 44: 73–80.
the Pacific islands for the biological control of insects and Pemberton, C. E., and J. S. Rosa. 1946. Life history of a new
other pests. So. Pacif. Comm. Tech. Pap. No. 101. parasite of the black widow spider in Hawaii. Hawaii. Plant-
Dzulhelmi, M. N., S. Suriyanti, M. Zulqarnain, and N. Che ers’ Rec. 50: 29–37.
Yusoff. 2014. New distributional records of spiders (Arach- Platnick, N. I. 2014. The world spider catalog, version 16.
nida: Araneae) from the west coast of Peninsular Malaysia. American Museum of Natural History. (http://www.wsc.
Pakistan J. Zool. 46: 1573–1584. nmbe.ch/genusdetail/3502, accessed August 2015).
Figueira, J.E.C., and J. Vasconcellos-Neto. 1993. Reproduc- Ramialiharisoa, A., L. de Haro, J. Jouglard, and M. Goyf-
tive success of Latrodectus geometricus (Theridiidae) on fon. 1994. Latrodectism in Madagascar. Med. Trop. (Mars)
Paepalanthus bromelioides (Eriocaulaceae): rosette size, 54: 127–130.
microclimate, and prey capture. Ecotropicos 5: 1–10. Reyes-Lugo, M., T. Sánchez, H. J. Finol, E. E. Sánchez, J.
Forster, R. R., and L. M. Forster. 1999. Spiders of New A. Suárez, B. Guerreiro, and A. Rodrı́guez-Acosta.
Zealand and their Worldwide Kin. Univ. of Otago Press, Dun- 2009. Neurotoxic activity and ultrastructural changes in mus-
edin, New Zealand. Dunedin, p. 270. cles caused by the brown widow spider Latrodectus geometri-
Fullaway, D. T. 1953. Three new species of Eurytoma (Hyme- cus venom. Rev. Inst. Med. Trop. Sao Paulo 51: 95–101.
noptera: Eurytomidae). Proc. Hawaiian Entomol. Soc. 15: Rinaldi, I.M.P. 2005. Spiders of a young plantation of eucalypt:
33–36. diversity and potential predator of the most frequent arboreal
Garb, J. E., A. Gonzalez, and R. G. Gillespie. 2004. The species. Acta Biol. Paranaense 34: 1–13.
black widow spider genus Latrodectus (Araneae: Theridii- Schreiner, I. 1989. Biological control introductions in the Caro-
dae): phylogeny, biogeography, and invasion history. Mol. line and Marshall Islands. Proc. Hawaii. Entomol. Soc. 29:
Phylogen. Evol. 31: 1127–1142. 57–69.
Goddard, J., S. Upshaw, D. Held, and K. Johnnson. 2008. Simó, M., M.F.R. Dias, C. Jorge, M. Castro, M. A. Dias, and
Severe reaction from envenomation by the brown widow spi- A. Laborda. 2013. Habitat, redescription and distribution of
der, Latrodectus geometricus (Araneae: Theridiidae). South Latrodectus geometricus in Uruguay (Araneae: Theridiidae).
Med. J. 101: 1269–1270. Biota Neotrop. 13: 371–375.
Hanson, P. E., and I. D. Gauld. 2006. Hymenoptera de la Suchard, J. R. 2009. Images in emergency medicine. Ann.
Region Neotropical. Mem. Am. Entomol. Inst. 77: 1–994. Emerg. Med. 54: 8–11.
Heeres, A. 1991. Natural history observations of the brown Triana, E., G. Barrantes, and P. Hanson. 2012.
widow spider Latrodectus geometricus (Araneae: Theridii- Incidence of parasitoids and predators on eggs of seven
dae). Naturalist 35: 31–34. species of Theridiidae (Araneae). Bull. Brit. Arachnol. Soc.
Herting, B. 1971. Arachnida to heteroptera: A catalogue of par- 15: 293–298.
asites and predators of terrestrial arthropods. Section A. Host Vetter, R. S. 2013. How to identify brown widow spiders,
or Prey/Enemy. Commonwealth Agricultural Bureaux, (http://cisr.ucr.edu/identifying_brown_widow_spiders.html,
Slough, England. accessed August 2015).
Kerr, A. M. 2013. Annotated checklist of the arachnids and myr- Vetter, R. S., L. S. Vincent, D.W.R. Danielsen, K. I. Rein-
iapods of the Mariana Islands, Micronesia, p. 16. ker, D. E. Clarke, A. A. Itnyre, J. N. Kabashima, and M.
Kiriakos, D., P. Núñez, Y. Parababire, M. Garcia, J. Med- K. Rust. 2012a. The prevalence of brown widow and black
ina, and L. D. Souza. 2008. First case of human latrodec- widow spiders (Araneae: Theridiidae) in urban southern Cali-
tism in Venezuela. Rev. Soc. Bras. Med. Trop. 41: 202–204. fornia. J. Med. Entomol. 49: 947–951.
8 JOURNAL OF MEDICAL ENTOMOLOGY
Vetter, R. S., L. S. Vincent, A. A. Itnyre, D. E. Clarke, K. I. Vincent, L. S., R. S. Vetter, W. J. Wrenn, J. K. Kempf, and J.
Reinker, D.W.R. Danielsen, L. J. Robinson, J. N. Kaba- E. Berrian. 2008. The brown widow spider Latrodectus geo-
shima, and M. K. Rust. 2012b. Predators and parasitoids of metricus C. L. Koch, 1841, in southern California. Pan-Pac.
egg sacs of the widow spiders, Latrodectus geometricus and Entomol. 84: 344–349.
Latrodectus hesperus (Araneae: Theridiidae), in southern
California. J. Arachnol. 40: 209–214. Received 2 June 2015; accepted 1 August 2015.