90 urn:lsid:arphahub.com:pub:4EBD0208-8328-5D69-8C44-EC50939C0967 urn:lsid:zoobank.org:pub:52E200AF-408A-423E-8797-5BFC4FDE0F99 Herpetozoa Herpetozoa 1013-4425 2682-955X Pensoft Publishers 10.3897/herpetozoa.39.e176733 176733 Review Article Amphibia Behaviour Asia World Consumption of the frog Hylarana nicobariensis Stoliczka, 1870 by a crab Parathelphusa pulcherrima (De Man, 1902), with a mini review of feeding on amphibians by crabs Quah Evan Seng Huat evanquah@ums.edu.my https://orcid.org/0000-0002-5357-1953 1 2 Conceptualization Writing - original draft Data curation Formal analysis Investigation Methodology Project administration Resources Supervision Validation Visualization Dukit Aderina https://orcid.org/0009-0003-3890-2560 3 Writing - review and editing Investigation Methodology Grismer Larry Lee https://orcid.org/0000-0001-8422-3698 4 5 6 Writing - review and editing Supervision Validation Hong Zijia https://orcid.org/0009-0005-4554-1315 7 Writing - review and editing Data curation Formal analysis Investigation Methodology Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400 Kota Kinabalu, Sabah, Malaysia Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah Kota Kinabalu Malaysia https://ror.org/040v70252 Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, Singapore 117377, Singapore Lee Kong Chian Natural History Museum, National University of Singapore Singapore Singapore School of Science, Monash University Malaysia, Bandar Sunway 47500, Malaysia School of Science, Monash University Malaysia Bandar Sunway Malaysia Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA Department of Biology, La Sierra University Riverside United States of America https://ror.org/05g1rjn35 Department of Herpetology, San Diego Natural History Museum, PO Box 121390, San Diego, California 92112, USA Department of Herpetology, San Diego Natural History Museum San Diego United States of America https://ror.org/00kmpab62 El Serpentario y C.E.M.A de Baja California Sur, Brercha California e/ Nuevo Reforma y Guaycura, Fracc: Benito Juárez, CP 23090, La Paz, Baja California Sur, Mexico El Serpentario y C.E.M.A de Baja California Sur La Paz Mexico School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia School of Biological Sciences, Universiti Sains Malaysia Penang Malaysia https://ror.org/02rgb2k63

Corresponding author: Evan Seng Huat Quah (evanquah@ums.edu.my)

Academic editor: Christoph Leeb

 2026 15 06 2026 39 253 260 ADAA1EBD-2EA5-5CB0-B5A7-4CB9E3E76EEC 10FFC552-0D5E-4D19-8586-EA1E03AB8939 01 11 2025 26 04 2026 This is an open access article distributed under the terms of the CC0 Public Domain Dedication. https://zoobank.org/10FFC552-0D5E-4D19-8586-EA1E03AB8939

We present a first record of feeding by the freshwater crab species, Parathelphusa pulcherrima on the frog species Hylarana nicobariensis (Ranidae) from Sabah (Borneo, Malaysia) together with a comprehensive review of crab predation on amphibians. A total of 32 crab species identifiable to 27 genera have been recorded to prey on frogs from at least 42 species in 35 genera, representing 21 families and one species of salamander to-date. The majority of the predation accounts were from the Neotropics and Afrotropics subcontinent.

 Borneo Crustacea Decapoda ecology Feeding behaviour frog Gecarcinucidae Malaysia natural history Introduction

Amphibians are important prey for many different predators in various ecosystems, including different species of macroinvertebrates such as crabs (McCormick and Polis 1982; Toledo 2005; Wells 2007; Valdez 2020). In tropical ecosystems, such as rainforests, where both groups thrive, predation by crustaceans, especially crabs of the infraorders Brachyura (true crabs) and Anomura (hermit crab and relatives) upon frogs is relatively common. In Brazil, for example, 10 frog species from three families have been recorded being preyed upon by six crab species (Affonso and Signorelli 2011; Caldart et al. 2011; Andrade et al. 2012; Marques et al. 2013; Motta-Tavares et al. 2015; Nogueira-Costa et al. 2016; Sichieri et al. 2021).

The island of Borneo in Sundaic Southeast Asia is well-known for hosting a high diversity of Amphibia with approximately 226 species known (Inger 1966; Inger et al. 2017; Haas et al. 2022; Bauer and Das 2024), but it is also rich in terrestrial and freshwater crab species with 106 species recorded (Grinang 2022). The number of species for both groups continues to increase as new species are continuously being discovered and described from different parts of Borneo (Ng 2021; Ng and Grinang 2022; Ng et al. 2022; Matsui et al. 2024; Dehling et al. 2025; Gonggoli et al. 2025a, 2025b; Imbun et al. 2025; Quah et al. 2025). Taxonomic effort aside, there have been no reports of crabs feeding upon any anurans on the island until now. Here, we describe a novel observation of a feeding interaction by a Bornean freshwater crab species, Parathelphusa pulcherrima (De Man, 1902), on a Cricket Frog, Hylarana nicobariensis Stoliczka, 1870, in Sabah. Hylarana nicobariensis is a common frog species that is widely distributed in the Indo-West Pacific, from the Nicobar Islands in the west, eastwards through southern Thailand and much of Sundaland including Peninsular Malaysia, Sumatra, Java, Borneo, Palawan and many of its surrounding islands (Malkmus et al. 2002; Inger et al. 2017; AmphibiaWeb 2025; Frost 2025). The species is a human commensal that thrives in anthropogenically modified landscapes, such as forest margins, logging roads, rural villages, roadside ditches and wet grassy areas. It is active both diurnally and nocturnally and breeds year-round whenever there are suitable environmental conditions in shallow ditches or temporary ponds (Malkmus et al. 2002; Inger et al. 2017; Haas et al. 2018). To contextualise the observation, we conducted a thorough literature review of published records of predation events by crabs on amphibians to assemble a dataset as a reference for future observations and research.

 Materials and methods Observation

As part of herpetofaunal surveys in Universiti Malaysia Sabah’s (UMS) main campus for a preliminary study of herpetofauna diversity in urban and suburban areas around Kota Kinabalu, Sabah, conducted by the first two authors (Evan S. H. Quah and Aderina Dukit), ecological/behavioural interactions observed opportunistically, such as the present feeding event, were recorded and photographed using an iPhone XS. The surrounding habitats and the weather conditions were also noted.

 Literature survey

The methodology used to conduct the literature survey was adapted from that employed by Quah et al. (2022, 2023), Quah and Chong (2023) and Quah and Grismer (2024). A comprehensive review of the published literature reporting predation by crabs on amphibians was conducted predominantly via online searches. Some of the literature was obtained via keyword searches in Google Scholar using combinations of the key words: crab, Anomura, Brachyura, Decapoda, eat, predation, prey on amphibians, Anura, frogs, salamanders, newts, caecilians. Other relevant work not available for downloading from this search were obtained manually by the authors by making library loan requests or by contacting the respective publication’s author(s) to obtain copies of their papers. The predatory crabs and their amphibian prey species recorded to the most specific taxonomic category possible, specific life stages of the amphibians (i.e. adult, juvenile, metamorph, tadpole and eggs) and the country of the observation were noted (Table 1). We follow the amphibian classification of Frost (2025) and taxonomy was updated from original reports with original taxon names indicated with superscripted text above the table. Literature was assessed up to 28 November 2025.

Compilation of amphibian prey species and known or suspected predatory crabs. a as Centrolenella granulosa; b as Limnonectes kuhlii; c as Scinax cf. arduous; d as Litoria aurea; e as Pelophylax cerigensis; M refers to marine crab species; W refers to wide-ranging crab species; * indicate laboratory observations; † indicate possible predation; Life stages are abbreviated as adult (A), juvenile (J), metamorph (M), tadpole (T) and eggs (E).

Crab Infraorder Brachyura
Prey Species Life Stage Crab Species Country Source
Salamanders
Salamandridae
Salamandrina terdigitata A †Potamon fluviatile Italy Rocca et al. (2005)
Frogs
Aromobatidae
Mannophryne olmonae A Eudaniela garmani Trinidad and Tobago Murphy (1997)
Mannophryne trinitatis A Unidentified Trinidad and Tobago Murphy (1997)
Bufonidae
Duttaphrynus melanostictus A †Balssipotamon fruhstorferi Vietnam Jestrzemski and Schütz (2016)
Rhinella ornata A Trichodactylus fluviatilis Brazil Segadilha and Silva-Soares (2015)
Centrolenidae
Cochranella granulosa a E Sesarma roberti Costa Rica Hayes (1983)
Conrauidae
Conraua alleni A Liberonautes latidactylus Guinea Schäfer et al. (2019)
Dendrobatidae
Colostethus inguinalis A Unidentified Panama Duellman and Trueb (1994)
Dendrobates auratus T *Armases angustumM Panama Gray and Christy (2000)
Dicroglossidae
Limnonectes fujianensis b A Candidiopotamon rathbuni Taiwan Tsuji (2005)
Limnonectes magnus A Isolapotamon mindanaoense Philippines Magdua et al. (2022)
Eleutherodactylidae
Eleutherodactylus cuneatus A Epilobocera cubensis Cuba Fong et al. (2012)
Hylidae
Boana raniceps A Dilocarcinus pagei Brazil Sichieri et al. (2021)
Bokermannohyla circumdata E Trichodactylus fluviatilis Brazil Nogueira-Costa et al. (2016)
Isthmohyla pseudopuma A Epibolocera sinuatifrons Costa Rica Warrington and Cossel (2012)
Ololygon cf. arduoa c A †Trichodactylus fluviatilis Brazil Marques et al. (2013)
Pseudis paradoxus A †Dilocarcinus pagei Brazil Affonso and Signorelli (2011)
Scinax acuminatus A Dilocarcinus pagei Brazil Sichieri et al. (2021)
Hylodidae
Crossodactylus gaudichaudii T †Trichodactylus petropolitanus Brazil Motta-Tavares et al. (2015)
Crossodactylus schmidti A Trichodactylus kensleyi Brazil Caldart et al. (2011)
Leptodactylidae
Leptodactylus latrans J Dilocarcinus pagei Brazil Affonso and Signorelli (2011)
Leptodactylus macrosternum A Goniopsis cruentata M Brazil Andrade et al. (2012)
Leptodactylus podicipinus A Dilocarcinus pagei Brazil Sichieri et al. (2021)
Leptodactylus cf. podicipinus J †Dilocarcinus pagei Brazil Affonso and Signorelli (2011)
Physalaemus pustulosus A Unidentified Trinidad and Tobago Murphy (1997)
Physalaemus pustulosus A Potamocarcinus richmondi Panama Ryan et al. (1981)
Mantellidae
Boophis rufioculis A Hydrothelphusa sp. Madagascar Rosa et al. (2014)
Mantidactylus cowanii A Unidentified Madagascar Abraham and Hutter (2022)
Megophryidae
Leptobrachium lumadorum T Isolapotamon sinuatifrons Philippines Lorenzo et al. (2017)
Microhylidae
Chiasmocleis ventrimaculata T Dilocarcinus sp.; Goyazama sp. Peru Schlüter and Salas (1991)
Chiasmocleis ventrimaculata A *Goyazama sp. Peru Schlüter and Salas (1991)
Hamptophryne boliviana T Dilocarcinus sp.; Goyazama sp. Peru Schlüter and Salas (1991)
Kaloula walteri A Sundathelphusa longipes Philippines Sy et al. (2015)
Odontobatrachidae
Odontobatrachus arndti A †Unidentified Guinea Schäfer et al. (2019)
Pelobatidae
Pelobates cultripes T Callinectes sapidus W Spain Alberto (2018)
Pelodryadidae
Ranoidea aurea d A, T Leptograpsus variegatus M Australia Pyke et al. (2013)
Phrynobatrachidae
Phrynobatrachus krefftii A Unidentified Tanzania Hirschmann and Hödl (2006)
Phyllomedusidae
Cruziohyla calcarifer A Ptychophallus uncinatus Costa Rica Wehrtmann et al. (2019)
Pipidae
Xenopus laevis A Potamonautus sp. South Africa Gutsche and Elepfandt (2007)
Xenopus mellotropicalis A Sudanonautes floweri Gabon Pollet et al. (2024)
Pyxicephalidae
Natalobatrachus bonebergi E Unidentified South Africa Acker-Cooper et al. (2025)
Ranidae
Aquarana catesbeiana J Parathelphusa reticulata Singapore Chen et al. (2025)
Hylarana nicobariensis A †Parathelphusa pulcherrima East Malaysia (Borneo) This study
Odorrana amamiensis A Eriocheir japonica W Japan Komine (2020)
Pelophylax ridibundus e A Potamon karpathos Greece Christopoulos et al. (2024)
Pelophylax sp. A Callinectes sapidus W Spain Pérez-Sorribes and Gil-Climent (2021)
Pelophylax sp. M Potamon fluviatile Italy Nicola et al. (2025)
Crab Infraorder Anomura
Unidentified frog
Unidentified frog A Birgus latro M - Helfman (1979)
Results Observation

On 7 March 2024 at approximately 17:30 h, a crab (ca. 40 mm carapace width) was observed feeding on an adult Hylarana nicobariensis of ca. 55 mm snout–vent length (Fig. 1), at Universiti Malaysia Sabah’s (UMS) main campus, specifically in a drain culvert opposite the Azman Hashim UMS Gallery (6.0418°N, 116.1146°E; alt. 55 m). The crab was tentatively identified as an adult specimen of Parathelphusa pulcherrima from the margin of its external orbital angle that is entire and smooth (Ng et al. 2022; Darren C.J. Yeo and Peter K.L. Ng in litt.).

10.3897/herpetozoa.39.e176733.figure1 2581B5BE-8165-534C-BC4C-C0DE4E233A09

The crab Parathelphusa pulcherrima feeding on Hylarana nicobariensis on UMS main campus in Kota Kinabalu, Sabah, East Malaysia, Borneo. Photos by Evan S.H. Quah.

https://binary.pensoft.net/fig/1681146

The prey frog was identified as H. nicobariensis, based on the following diagnostic characteristics: body slender; head long, narrow; snout pointed, projecting beyond lower jaw; tympanum large and distinct; long, slender hind limbs; skin of dorsum finely pebbled; continuous thin dorsolateral fold extends from eye to pelvic region; colouration of dorsum brown with dark spots and marbling; hind-limbs brown with darker spots and barring; black streak on side of head beginning from the snout, along canthus rostralis continuing posterior to eye and covering the tympanum on to the flanks at the insertion point of the fore-limb; upper lips with a uniform white streak extending to the rictal ridge behind the jaw (Malkmus et al. 2002; Inger et al. 2017; Haas et al. 2018; Fig. 2).

10.3897/herpetozoa.39.e176733.figure2 DC546971-408C-5B7D-8E05-DD9F1EC789BE

Typical-looking Cricket Frog (Hylarana nicobariensis) found in UMS campus. Photo by Evan S.H. Quah.

https://binary.pensoft.net/fig/1681147

The observation took place in a drain culvert that had numerous puddles of water that formed in cracks and eroded parts of the concrete floor. Rain the previous day raised the water level to relatively high (3–5 cm depth in different spots), while it was overcast during the time of the observation. When we chanced upon the event, the crab was already feeding on the frog’s carcass at a depth of approximately 4 cm of water. It cannot be determined whether this was a result of predation or a scavenging event, in which the crab was feeding upon an already dead frog. The crab had grasped the frog by its right fore-limb, sheering away the flesh with its mouthparts. The skin had been torn off and the muscles and bone exposed on the right forelimb. At the time of the observation, the crab had already eviscerated the gut of the frog and all the toes and much of the left rear limb of the frog had been eaten (Fig. 1A). We continued our observation for 10 min, during which the crab retreated slightly into a crevice in the concrete still hanging on to the fore-limb. We returned to observe the crab 15 min later after completing surveys of a nearby area and it had returned to feeding on the frog in the open and had started to feed on the right hind-limb (Fig. 1B). Another specimen of H. nicobariensis was observed in the same culvert during the predation event and other herpetofaunal species observed were an undetermined species of the Limnonectes kuhlii (Tschudi) group, Gekko monarchus (Schlegel) and Eutropis rudis (Boulenger).

 Crab predation on amphibians

A total of 32 crab species that were identifiable to at least genus level representing 27 genera have been recorded to predate upon frogs from at least 42 species in 35 genera of 21 families and one species of salamander (Table 1). No reports of species of caecilians falling prey to crabs were recorded in our review of published literature. The highest number of frog prey species belong to the family Hylidae with six species recorded, whereas the main predatory crab species recorded were Trichodactylus fluviatilis Latreille and Dilocarcinus pagei Stimpson both of the family Trichodactylidae and infraorder Brachyura (Crab Database 2025). The majority of the predation accounts were from the Neotropics (Trinidad and Tobago, Brazil, Costa Rica, Panama and Peru) and the Afrotropics (Guinea, Tanzania, South Africa, Gabon and Madagascar). There are no known predation events reported from North America (Table 1).

 Discussion

This report of a Parathelphusa pulcherrima crab feeding on Hylarana nicobariensis is the first for a feeding interaction involving these two species. It also increases the record for the number of frog species from the family Ranidae known to be fed upon by crabs which now includes H. nicobariensis, Aquarana catesbeiana (Shaw), Odorrana amamiensis (Matsui), Pelophylax ridibundus (Pallas) and Pelophylax sp. (Table 1), (Komine 2020; Pérez-Sorribes and Gil-Climent 2021; Christopoulos et al. 2024; Nicola et al. 2025). It is also the first time a member of the genus Hylarana has been reported as a food/potential prey item for crabs.

The collated data also indicate that crabs are a significant arthropod predator of frogs and this behaviour is taxonomically and geographically widespread. In their early review, McCormick and Polis (1982) listed two reports of predation events by crabs, one by an unidentified crab species preying on tadpoles of an unidentified frog and the second was that by a Coconut Crab (Birgus latro) also on an unidentified frog (Wager 1965; Helfman 1979). Subsequently, Toledo (2005) also reported two species of frogs falling prey to crabs. The first cited the predation record of B. latro reported by Helfman (1979) and the second species was based on Duellman and Trueb (1994) that recorded Colostethus inguinalis being preyed upon by an unidentified freshwater crab. Our extensive review substantially increased the number of records (Table 1). Our reported observation is also the second time a crab of the genus Parathelphusa has been recorded feeding on an amphibian, with the first being a very recent record of P. reticulata feeding on Aquarana catesbeiana in Singapore (Chen et al. 2025). Our data compilation also indicates that the majority (84.4%: 27 out of the 32 recorded species) of the crab predators are freshwater or terrestrial species which is understandable given that there are so few species of saline-tolerant frogs (Table 1). Finally, the assembled data demonstrate that crabs are known to prey on all developmental stages of amphibians (Table 1).

Crabs feeding on amphibians have now been reported from all continents, except for North America and Antarctica (Table 1). No recent amphibians occur in Antarctica given its perpetually frigid conditions, therefore predation events are impossible. The cause for the lack of observations in North America is uncertain, but it could be the lower diversity of freshwater and terrestrial crab species further north of the Equator. Our new observation also constitutes the first report of batrachophagy by a crab for Malaysia from the island of Borneo. Studies in other areas have demonstrated that crabs may be the predominant predator of frogs living along small, forest streams and interactions between frogs and crabs are probably more common than is presently known as many taxa are semi-aquatic and syntopic in many habitats (Tsuji 2005). Crabs are also notable predators of many other species of herpetofauna as well, such as turtles and snakes (McCormick and Polis 1982; Dehling 2008; Jablonski et al. 2024). Likewise, other crustaceans are also known to prey upon amphibians, such as crayfish of the infraorder Astacidea (Axelsson et al. 1997; Wilson and Williams 2014; Ott 2016; Stawikowski 2021) and copepods that have been observed preying on larval salamanders of the genus Ambystoma (McCormick and Polis 1982). Through more extensive fieldwork, interactions between additional crustacean and herpetofauna species will likely be observed and recorded in the future.

 Acknowledgements

Our team is grateful to the Sabah Biodiversity Centre (SaBC) for granting us permits to conduct our research [JKM/MBS.1000-2/2 JLD 14 (19), JKM/MBS.1000-2/2 JLD 17 (77), JKM/MBS.1000-2/2/1 JLD 1 (30), JKM/MBS.1000-2/2/1 JLD 1 (33) and JKM/MBS.1000-2/2 JLD 15 (1)]. This research was partially funded by the Nagao Natural Environment Foundation (NEF) and is part of a preliminary study into the diversity of herpetofauna in urban and suburban areas around Kota Kinabalu, Sabah (UMS project code: LPA2207). This project was approved by the animal ethics committee of UMS (UMS/IBTP7.2/800-2/1/17 [file no.: AEC 0011/2022]). We also thank Darren C.J. Yeo and Peter K.L. Ng of the Lee Kong Chian Natural History Museum, Singapore for helping us identify the crab.

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