Herpetology Notes, volume 14: 291-296 (2021) (published online on 09 February 2021)

Predation by a Brown Widow Spider, Latrodectus geometricus (Koch, 1841), on a Common Dwarf Gecko, Lygodactylus capensis (Smith, 1849), with a review of the herpetofaunal diet of Latrodectus spiders

Daniel van Blerk1, John Measey1,*, and James Baxter-Gilbert1

The Common Dwarf Gecko, Lygodactylus capensis box (Fig. 1A). The lizard was initially found at 17:13 h, (Smith, 1849), is a small, diurnal gekkonid with a writhing in the web and suspended above its autotomised wide African distribution, ranging from Kenya to tail, which lay on the base of the electrical box. At this northeastern South Africa and extending westward time, the spider was present on the electrical box, but not into Namibia and Angola (Rebelo et al., 2019). This in contact with the lizard. Shortly after being observed species is naturally absent from western South Africa, and identified, the spider retreated into the refuge of the but invasive populations have been observed scattered electrical box. By 17:18 h the lizard was motionless and across much of the country – including in large urban appeared dead, its hind legs having been wrapped against centres such as Bloemfontein, Cape Town, East London, the body with spider silk (Fig. 1B). On the final visit to George, Makhanda (formerly Grahamstown), and Port the electrical box at 17:48 h, the deceased lizard had been Elizabeth (Rebelo et al., 2019; Conradie et al., 2020). removed from its original position in the web and could The successful spread of L. capensis may be attributed not be seen. We presume the spider had pulled the body to this lizard’s proclivity for urban ecosystems and other further into its refuge to feed on it safely without further heavily human-modified habitats, as these lizards are able disturbance. to exploit novel resources and can capitalise on human Previous reports have stated that these lizards are transportation networks to facilitate extralimital range frequent prey items for a variety of vertebrates, including expansions (Rebelo et al., 2019). This anthropophilic birds, lizards, mammals, and snakes (Simbotwe, 1983; behaviour, however, may also result in L. capensis Maritz and Maritz, 2020a,b), as well as predatory encountering other urbanised taxa more frequently – invertebrates, including Rain Spiders (genus Palystes), including potential predators such as Latrodectus spiders. Baboon Spiders (genus Harpactirella), Crab Spiders Here we report one such encounter, during which a L. (genus Misumena), and mantids (genus Sphodromantis) capensis was seen to have been attacked by a Brown (Warren, 1923; Bauer, 1990; Maritz and Maritz, 2020a,b). Widow Spider, Latrodectus geometricus (Koch, 1841), To the best of our knowledge, this is the first observation in one of the lizard’s invasive populations near George, of a predation event by L. geometricus on L. capensis. South Africa (centred on 33.99°S, 22.45°E). Latrodectus geometricus has a near-global distribution On 19 August 2020 at a private residence in Heather related to its anthropophilic nature, with known invasive Park, George, Western Cape, South Africa (33.9581°S, populations in North and South America, Hawaii, and 22.4213°E), the first author observed an adult dwarf across Australasia (Taucare-Ríos et al., 2016), and it gecko (L. capensis) entangled within a brown widow (L. is presumed to be native to southern Africa (see Garb geometricus) web inside an outdoor dual-socket electrical et al., 2004; Brown et al., 2008; Measey et al., 2020). Across its native and invasive range, L. geometricus has been seen to feed on a wide variety of herpetofauna (Table 1). Known African reptile prey items in the 1 Centre for Invasion Biology, Department of Botany and native range include Herald Snakes, Crotaphopeltis Zoology, Stellenbosch University, Stellenbosch, Western hotamboeia (Laurenti, 1768), Brown House Snakes, Cape, South Africa. Boaedon capensis (Boie, 1827), and Kalahari Tree * Corresponding author. Email: [email protected] Skinks, Trachylepis spilogaster (Peters, 1882) (Bayliss, © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. 2001; Maritz and Maritz 2020a,b). Observations of 292 Daniel van Blerk et al.

Figure 1. A live adult Common Dwarf Gecko, Lygodactylus capensis, entangled in the web of a Brown Widow Spider, Latrodectus geometricus, at Heather Park, George, Western Cape, South Africa. The lizard was alive at 17:13 h (A) and appeared to be dead by 17:18 h (B). Upon first observation, the spider was seen and identified, however it retreated before it could be photographed and remained reluctant to emerge. The autotomised tail can be seen suspended in the web between the switches as indicated by the red arrow in (A).

L. geometricus feeding on herpetofauna within their frenatus in Mexico (de Luna et al., 2020) or L. capensis. invasive ranges in the Americas include Scarlet Snakes, The increasing expansion of urban areas globally Cemophora coccinea (Blumenbach, 1788) (Stevenson (Chen et al., 2020), along with continued transport and and Crook, 2018), Asian House Geckos, Hemidactylus propagation of invasive species (Seebens et al., 2017), have frenatus Duméril & Bibron, 1836 (de Luna et al., 2020), led to novel predator-prey interactions in anthropogenic Brahminy Blindsnakes, Indotyphlops braminus (Daudin, habitats. Although in general both L. capensis and L. 1803) (Martínez-Coronel and Navarrete-Jiménez, 2018), geometricus are native southern African species, the fact and Black-headed Snakes, Tantilla melanocephala that they both capitalise on human habitats may result in (Linneaus, 1758) (Rocha et al., 2017). Taken together, increased interactions between them, especially as urban it is clear that these spiders are formidable predators of populations in their native and invasive ranges expand. small-bodied reptiles. Overall, this observation adds both to the growing list of In general, spiders of the genus Latrodectus have known predators for L. capensis and known vertebrate a global reputation for feeding on vertebrate prey, prey for Latrodectus spiders. including a variety of herpetofauna – encompassing frogs, lizards, and snakes (Table 1). As a group, these Acknowledgments. We thank Robin and Bryan Maritz for their insight into the variety of known predators of L. capensis and spiders possess potent venom and strong webs, enabling access to their Predation Records Online Data Repository, which them to capture and kill prey items considerably larger was immensely helpful in writing this manuscript. We would also than themselves (O’Shea and Kelly, 2017; Nyffeler and like to thank the DSI-NRF Centre of Excellence for Invasion Vetter, 2018). Furthermore, Latrodectus spiders have a Biology for their support. tendency to thrive in anthropogenic habitats, possibly associated with beneficial structural, thermal, and hydric References conditions associated with urban landscapes, which has Arteaga-Tinoco, I., Anguiano-Peña, J., Raya-García E., Alvarado- further expanded their cosmopolitan distribution (Vink et Díaz, J. (2016): Natural history notes. Hyla eximia (Mountain al., 2011). Thus, it is unsurprising that they have been Tree Frog). Predation. Herpetological Review 47: 641–642. regularly seen feeding on other anthropophilic taxa, such Bayliss, P.S. (2001): Natural history notes. Lamprophis fuliginosus as invasive and human-commensal herpetofauna, like H. (Brown House Snake). Predation. Herpetological Review 32: 48–49. Latrodectus Predation on Amphibians and Reptiles 293

2002

2013 1990 Piñero, 2002 Piñero, 2002 Piñero, 2002 Piñero ,

- - - - Baurecht, 1988

Werner, 1989 , Ridgeway , nchez nchez nchez nchez and Kamees, 2010 Kamees, 2013 Feldman, 2019 Tinoco et al., 2016 2007 - and 1990 , , 1973; , Raven , , and dar Sá and dar Sá and dar Sá and dar Sá and Zilberberg, 1988 Schwammer Hó Blondheim and Hó Painter Hó Reference Arteaga Thill König, 1987; Orange, 2007 Wilson, 1991; Lane, 2012 Orange Copland, 1953 1943 McKeown, Orange, 2007 de Luna et al., 2020 Hó Raven Metcalfe and , , Colombo , Cook

Species Chamberlin & Ivie, 1935 Ivie, & Chamberlin Fabricius, 1775

Charitonov, 1954 Thorell, 1870 Melic, 2000

geometricus tredecimguttatus lilianae L. L. L. revivensis Shulov, 1948 Rossi, 1790 tredecimguttatus L. lilianae L. L. pallidus pallidus L. Latrodectus L. mactans hasselti L. lilianae L. L. hesperus L. mactans hasselti L. hasselti L. hasselti L. L. hesperus L. hesperus hasselti L. lilianae L. hasselti L. hasselti L. L. hasselti L. L.

(Schinz, 1833) Fitzinger, 1826 spiders. Latrodectus (Duméril Bibron,& 1839) (Sternfeld, 1918) (Duméril Bibron,& 1839) Girard, 1858 (Linnaeus, 1758) Duméril, 1851 (Gray, 1845) (Linnaeus, 1758) (De Vis, 1888) (Duméril Bibron,& 1836) (Braun, 1877) (Blainville, 1835) (Blainville,

(Baird, 1854) (Lichtenstein, 1823)

(Duméril Bibron,& 1836) Baird & Girard, 1852 Girard, & Baird

a a

a guichenoti

sp.

Podarcis melisellensis algirusPsammodromus Mesalina guttulat Mesalina Hylidae Anguidae Gekkonidae Lacertidae Phrynosomatidae Phyllodactylidae Scincidae Psammodromus hispanicusPsammodromus eximius Dryophytes multicarinata Elgaria marmoratus Christinus variegat Gehyra Acanthodactylus erythrurus guttulata Mesalina Phrynosoma hernandesi Uta stansburiana Tarentola mauritanica Anomalopus verreauxii Lampropholis delicata Lygosoma Hemidactylus frenatus frenatus Hemidactylus Lampropholis Chalcides ocellatus (Forskal, ocellatus Chalcides 1775) Cryptoblepharus pulcher Hemiergis quadrilineata

IZARDS ROGS

Prey Prey Taxon F L

Amphibian and reptile prey of 1. Table 294 Daniel van Blerk et al. 2018 2020a,b , Jiménez , , ,

Maritz 2014 1982 - Navarrete 2018 , , 2014 , , 2014 2014 , , al. et 2007

, , , , , ,

Polis , Crook ,

2017 Tucker Kelly Kelly Kelly, 2014 2011 , , Maritz 2020a,b Maritz 2020a,b

, ,

Carroll Coronel and Márquez 1990 - , , 1948 1948 Shea and Shea and Shea and Jones et al. Stevenson and and Beaman O’ Bayliss, 2001; Maritz and Maritz 2001; Bayliss, O’ Martínez - Ervin and Orange

Rocha et al. Rocha 1948 Neill, O’ Carbajal Bayliss, 2001 Reference , , Neill McCormick and McCormick Maritz and , , Neill Maritz and de Rebeira, 1981

Cambridge, 1904 -

Pickard

Species esperus h L. mactans hasselti L. hasselti L. L. L. geometricus L. geometricus L. mactans hasselti L.

L. mactans L. geometricus Latrodectus hasselti L. L. mactans L. mactans hasselti L. L. geometricus L. geometricus L. indistinctus L. L. mactans hasselti L. L. geometricus

(Laurenti, 1768) (Smith, 1849) (Shaw, 1794) (O’Shaughnessy, 1874) Cope Cope 1860 (Peters, (Peters, 1882) (Daudin, 1803) (Harlan, 1827) Boie, 1827) (Blumenbach, 1788) ( (Baird Girard,& 1852) Günther, 1872 (Say, 1822)

(Holbrook, 1839) (Storr, 1964)

(Günther, 1863) (Baird Girard,& 1852)

(Worrell, 1956)

(cont.)

Suta dwyeri tenuis Contia Crotaphopeltis hotamboeia Hypsiglena chlorophaea Suta nigriceps Scincidae Scincella lateralis Saproscincus mustelinus Aspidoscelis gularis Cemophora coccinea (Kennicott, euryxanthus 1860) Micruroides Boaedon fuliginosus Indotyphlops braminus Storeria dekayi Liochlorophis vernalis Suta monachus Pseudechis porphyriacus Teiidae Colubridae Elapidae Lamprophiidae Typhlopidae Tantilla melanocephala (Linnaeus, 1758) Pseudonaja affinis Trachylepis punctatissima spilogaster Trachylepis

(cont.) NAKES

Prey Prey Taxon S

Table 1 Table Latrodectus Predation on Amphibians and Reptiles 295

Bauer, A.M. (1990): Gekkonid lizards as prey of invertebrates and Maritz, R.A., Maritz, B. (2020b): Data for “Sharing for science: predators of vertebrates. Herpetological Review 21: 83–87. High-resolution trophic interactions revealed rapidly by social Beaman, K.R., Tucker, N.G. (2014): Natural history notes. media”. Figshare Dataset. Accessible at https://doi.org/10.6084/ Contiatenuis (Sharp-tailed Snake). Predation. Herpetological m9.figshare.11920128.v2. Accessed on 2 September 2020. Review 45: 514. Martínez-Coronel, M., Navarrete-Jiménez, A. (2018): Natural Blondheim, S., Werner, Y.L. (1998): Lizard predation by widow history notes. Indotyphlops braminus (Brahminy Blindsnake). spiders. The Herpetological Bulletin 30: 26–28. Spider predation. Herpetological Review 49: 754. Brown, K.S., Necaise, J.S., Goddard, J. (2008): Additions to the McCormick, S., Polis. G.A. (1982): Arthropods that prey on known U.S. distribution of Latrodectus geometricus (Araneae: vertebrates. Biological Reviews 57: 29–58. Theridiidae). Journal of Medical Entomology 45: 959–962. McKeown, K.C. (1943): Vertebrates captured by Australian Carbajal-Márquez, R.A., González-Saucedo, Z.Y., Quintero-Díaz, spiders. Proceedings of the Royal Zoological Society of New G.E. (2014): Natural history notes. Aspidoscelis gularis (Spotted South Wales 63: 16–30. Whiptail). Predation. Herpetological Review 44: 505. Measey, J., Robinson, T.B., Kruger, N., Zengeya, T.A., Hurley, B.P. Chen, G., Li, X., Liu, X., Chen, Y., Liang, X., Leng, J., et al. (2020): (2020): South Africa as a donor of alien animals. In: Biological Global projections of future urban land expansion under shared Invasions in South Africa, p. 787–830. van Wilgen, B., Measey, socioeconomic pathways. Nature Communications 11: 1–12. J., Richardson, D., Wilson, J., Zengeya, T., Eds., Cham, Colombo, M. (2013): Natural history notes. Chalcides ocellatus Switzerland, Springer. (Ocellated Skink). Spider predation. Herpetological Review 44: Metcalfe, D.C., Ridgeway, P.A. (2013): A case of web entanglement 320–321. and apparent predation of the skink Lampropholis delicata (De Conradie, W., Nicolau, G.K., Kemp, L., Edwards, S., Klien Vis, 1888) by the red-back spider Latrodectus hasseltii (Thorell, Snakenborg, S., Reeves, B. (2020): Lygodactylus capensis 1870) in an autochthonous mesic habitat in coastal southeast (Smith, 1849) Common Dwarf Gecko: New records from the Australia. Herpetology Notes 6: 375–377. Makhanda Region. African Herp News 74: 78–82. Neill, W.T. (1948): Spiders preying on reptiles and amphibians. Cook, R. (1973): The wall lizard Cryptoblepharus boutonii Herpetofauna 19(1): 31–32. virgatus. Herpetofauna 6(2): 15–16. Nyffeler, M., Vetter, R.S. (2018): Black widow spiders, Latrodectus Copland, S.J. (1953): Presidential address, delivered at the Seventy- spp. (Araneae: Theridiidae), and other spiders feeding on eighth Annual General Meeting, 25th March 1953, Recent mammals. The Journal of Arachnology 46: 541–548. Australian herpetology. Proceedings of the Linnean Society of O’Shea, M., Kelly, K. (2017): Predation on a Weasel Skink New South Wales 78: i–xxxvii. (Saproscincus mustelinus) (Squamata: Scincidae: Lygosominae) de Luna, M., Rodriguez, M., Barrios, R.G. (2020): Predation on the by a Redback Spider (Latrodectus hasselti) (Araneae: common house gecko Hemidactylus frenatus Schlegel, 1836 by Araneomorpha: Theridiidae), with a review of other Latrodectus the brown widow spider Latrodectus geometricus Koch, 1941 in predation events involving squamates. Herpetofauna 44: 49–55. Colima, Mexico. Herpetology Notes 13: 555–556. Orange, P. (1990): Predation on Rhinoplocephalus monachus de Rebeira, P. (1981): A redback spider attacking an immature (Serpentes: Elapidae) by the redback spider, Latrodectus dugite. Western Australian Naturalist 15: 33–34. mactans. Herpetofauna 20: 34. Ervin, E.L., Carroll, S.L. (2007): Natural history notes. Hypsiglena Orange, P. (2007): Predation on lizards by the red-back spider, torquata chlorophaea (Sonoran Nightsnake). Predation. Latrodectus hasselti. Herpetofauna 37: 32–35. Herpetological Review 38: 468. Painter, C.W., Kamees, L.K. (2010): Natural history notes. Garb, J.E., Gonzalez, A., Gillespie, R.G. (2004): The black widow Phrynosoma hernandesi (Greater Short-horned Lizard). spider genus Latrodectus (Araneae: Theridiidae): phylogeny, Predation. Herpetological Review 41: 227. biogeography, and invasion history. Molecular Phylogenetics Raven, R. (1990): Spider predators of reptiles and amphibia. and Evolution 31: 1127–1142. Memoirs of the Queensland Museum 29: 448. Hódar, J.A., Sánchez-Piñero, F. (2002): Feeding habits of the black Rebelo, A.M., Bates, M.F., Burger, M., Branch, W.R., Conradie, widow spider Latrodectus lilianae (Aranae: Theridiidae) in an W. (2019): Range expansion of the Common Dwarf Gecko, arid zone of south-east Spain. Journal of Zoology 257: 101–109. Lygodactylus capensis: South Africa’s most successful reptile Jones, L.L., King, A.D., Simpson, P.A., Taiz, J., Wolterbeek, invader. Herpetology Notes 12: 643–650. P. (2011): Natural history notes. Micruroides euryxanthus Rocha, C.R., Motta, P.C., de Souza Portella, A., Saboya, (Sonoran Coralsnake). Predation. Herpetological Review 42: M., Brandão, R. (2017): Predation of the snake Tantilla 440–441. melanocephala (Squamata: Colubridae) by the spider König, R. (1987): Die Schwarze Witwe (Latrodectus mactans Latrodectus geometricus (Araneae: Theridiidae) in Central hasselti Thorell) als Fressfeind von Reptilien in Australien. Brazil. Herpetology Notes 10: 647–650. Herpetofauna (Weinstadt) 9(48): 6–8. Schwammer, H., Baurecht, D. (1988): Der Karstläufer, Lane, J.S. (2012): Natural history notes. Uta stansburiana Podarcis melisellensis fiumana (Werner, 1891), als Beute (Common Side-Blotched Lizard). Mortality. Herpetological der Europäischen Schwarzen Witwe, Latrodectus mactans Review 43: 339 tredecimguttatus (Rossi, 1790). Herpetozoa 1: 73–76. Maritz, R.A., Maritz, B. (2020a): Sharing for science: high- Seebens, H., Blackburn, T.M., Dyer, E.E., Genovesi, P., Hulme, P.E., resolution trophic interactions revealed rapidly by social media. Jeschke, J.M. et al. (2017): No saturation in the accumulation of PeerJ 8: e9485. alien species worldwide. Nature Communications 8: 14435. 296 Daniel van Blerk et al.

Simbotwe, M.P. (1983): On spacing patterns and diversity of escape tactics in diurnal geckos (Lygodactylus) in Kafue Flats, Zambia. Amphibia-Reptilia 4: 35–41 Stevenson, D.J., Crook, C.W. (2018): Natural history notes. Cemophora coccinea (Scarletsnake). Predation. Herpetological Review 49: 543–544. Thill, V.L., Feldman, C. (2019): Understanding patterns of resistance to spider venom in lizards: ecology and phylogeny matter. Herpetological Review 50: 573. Taucare-Ríos, A., Bizama, G., Bustamante, R.O. (2016): Using global and regional species distribution models (SDM) to infer the invasive stage of Latrodectus geometricus (Araneae: Theridiidae) in the Americas. Environmental Entomology 45: 1379–1385. Vink, C.J., Derraik, J.G., Phillips, C.B., Sirvid, P.J. (2011): The invasive Australian redback spider, Latrodectus hasseltii Thorell 1870 (Araneae: Theridiidae): current and potential distributions, and likely impacts. Biological Invasions 13: 1003–1019. Warren, E. (1923): Lizard-eating S. African spider. Annals of the Natal Museum 5: 95–100. Wilson, B.S. (1991): Latitudinal variation in activity season mortality rates of the lizard Uta stansburiana. Ecological Monographs 61: 393–414. Zilberberg, G. (1988): Behavioural ecology of the widow spider Latrodectus revivensis. Shappirit 6: 52–77.

Accepted by Darren Pietersen