Use the following type of citation: North-western Journal of Zoology 2021: e211508
Paper Submitted to The North-Western Journal of Zoology
1 *Handling editor: Yurii Kornilev
2 *Manuscript Domain: Herpetology of the Americas
3 *Manuscript code: NwJZ_20_HA_12
4 *Submission date: 04_06_2020
5 *Revised: 15_03_2021
6 *Accepted: 23_03_2021
7 *No. of words (without abstract, acknowledgement, references, tables, captions): 2340 8 (papers under 700 words are not accepted)
9 *Editors only: 10
11 Zoology 12 Title of the paper: of
13 Diet and clutch size of Rhinella castaneotica (Anura: Bufonidae) from a forest area in Serra
14 do Navio, Amapá, Brazil Journal 15 Running head:
16 Diet and clutch size of Rhinella castaneotica paper 17 Authors (First LAST - without institution name!):
18 Vinícius Figueiredo, Edgar Santos, Fillipe Santos, Patrick Sanches, Carlos Campos North-Western 19 accepted
20 Key Words (at least five keywords):
21 Prey selection, trophic niche, niche breadth, reproduction, Brazil.
22
23 No. of Tables: 1
24 No. of Figures: 0
25 No. of Files (landscape tables should be in separate file): 1
26
27 Use the following type of citation: North-western Journal of Zoology 2021: e211508
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28 Diet and clutch size of Rhinella castaneotica (Anura: Bufonidae) from a forest area in
29 Serra do Navio, Amapá, Brazil
30 Vinícius A. M. B. de FIGUEIREDO*, Edgar Matos dos SANTOS, Fillipe PEDROSO-
31 SANTOS, Patrick R. SANCHES and Carlos Eduardo COSTA-CAMPOS
32 Universidade Federal do Amapá, Departamento de Ciências Biológicas e da Saúde,
33 Laboratório de Herpetologia, Campus Marco Zero do Equador, 68.903-419, Macapá, AP,
34 Brazil
35 * Corresponding author, V. A. M. B. de Figueiredo, E-mail: [email protected]
36
37 Abstract. We studied the diet and the clutch size of Rhinella castaneotica from a primary Zoology 38 forest in northern Brazil, municipality of Serra do Navio,of Amapá state. A total of 100 toads 39 (77 males and 23 females) were collected between April 2017 and March 2018 through active
40 searches. Thirty-four individuals (34%) had a total of 870 prey items in their stomach Journal 41 contents. Hymenoptera and Coleoptera were the most important and numerous prey items in
42 the diets of males and females, respectively. The niche breadth based on prey numbers was
43 0.24. Based on 9 females with eggs, thepaper number of ovarian eggs ranged from 199 to 720 eggs.
44 We conclude that R. castaneotica has a specialized diet and its clutch size is independent of
45 the female SVL andNorth-Western body mass. We highlight the importance of further studies that contribute accepted 46 to a better understanding of the conservation status and ecological aspects of such poorly
47 known anurans in the Amazon.
48
49 Key Words: prey selection, trophic niche, niche breadth, reproduction, Brazil.
50 Running title: Diet and clutch size of Rhinella castaneotica
51
52 Introduction Use the following type of citation: North-western Journal of Zoology 2021: e211508
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53 The natural history of a given species is characterized by its distinct life-history
54 components (Putnam 1994). Diet, reproduction and micro-habitat use make up the
55 components most often examined in the majority of studies regarding the natural history of
56 anurans species (De-Carvalho et al. 2008, Freitas et al. 2008, Luria-Manzano & Gutierrez-
57 Mayen 2014, Jorge et al. 2015, Pinto & Menin 2017).
58 As pointed by Toft (1981), several factors influence the diet of anurans, such as the
59 size of prey, foraging strategies, body size, and physiological constraints. Furthermore, the
60 diet may vary ontogenetically and due to sexual differences, as males and females have
61 different energy needs (Lima 1998, Wu et al. 2005). As result, there is plasticity involving
62 anurans within trophic chains, which is expressed by their acting as predators and prey Zoology 63 (Toledo et al. 2007). of 64 The genus Rhinella is currently composed of 92 species recognized, distributed from
65 southern Texas, USA to southern South America (Frost 2020). Species often show sexual Journal 66 dimorphism, as recorded in several species such as R. achalensis, R. marina, R. rubescens,
67 and R. diptycha (Monnet & Cherry 2002, Jofré et al. 2005, Arantes et al. 2015). In general,
68 the toads of the genus Rhinella exhibitpaper explosive breeding events (Machado & Bernarde
69 2011), and these reproductive behaviors have been well-documented (Pombal-Junior &
70 Haddad 2005, Vargas-SalinasNorth-Western 2007, Machado & Bernarde 2011, Bowcock et al. 2013, accepted 71 Valentim et al. 2013, Costa-Campos et al. 2016).
72 The R. margaritifera species group currently contains 14 species, including the poorly
73 known R. castaneotica (Caldwell 1991). This small-sized species (snout-vent length 18.4–
74 23.6 mm in males and 18.9–26.3 mm in females) is present in the Amazon basin of Bolivia,
75 Brazil, Colombia, and Peru (IUCN 2015). Rhinella castaneotica are nocturnal and terrestrial
76 toads; south of the Amazon River males call sporadically throughout the rainy season in Use the following type of citation: North-western Journal of Zoology 2021: e211508
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77 ponds and small holes filled with water or from within empty Brazil nut capsules called
78 “castanheiras” (Caldwell 1993, Lehtinen et al. 2004).
79 Herein, we provide data on some aspects of the natural history of R. castaneotica, including
80 information on the diet composition and reproduction in a population from a forest area in
81 Amapá State, Brazilian Amazonia.
82
83 Material and Methods
84 We conducted surveys at the Cancão Municipal Natural Park (0°54’39.04”N,
85 52°0’29.72”W), Serra do Navio municipality, Amapá state, Brazil. The region is composed of
86 a mosaic of seasonal ombrophilous dense forest and floodplain (igapó) forest (Drummond et Zoology 87 al. 2008). The climate is Am (Köppen-Geiger classification;of tropical monsoon, without a dry 88 season), and the mean annual rainfall is 1,885 mm and temperature 27 °C (Peel et al. 2007).
89 Individuals examined in this study were collected for a study of the natural history of Journal 90 anurans in the eastern Amazon (diet, reproduction and helminth parasites) using visual and
91 auditory active searches (Heyer et al. 1994) in temporary ponds and on the forest floor during
92 five consecutive nights monthly frompaper April 2017 to March 2018. Immediately after
93 collection, all individuals were weighed with a digital balance (0.1 g precision), killed using
94 lidocaine 2%, and theirNorth-Western snout-vent length (SVL) was measured with a digital caliper (0.01 accepted 95 mm precision). The sex was recorded for each individual based on direct observation of
96 gonads.
97 Individuals were fixed in 10% formaldehyde solution, preserved in 70% ethanol, and
98 housed in the Herpetological Collection of the Universidade Federal do Amapá, state of
99 Amapá, Brazil (vouchers CECC 1419, 1420, 1449, 1450, 1452–1463, 1480–1484, 1527–
100 1529, 1854–1866, 1872, 1873, 1875–1880, 1882–1888, 2020, 2063–2079, 2092–2099, 2126–
101 2135, 2142–2155, 2259–2262). Use the following type of citation: North-western Journal of Zoology 2021: e211508
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102 For dietary analysis we removed the stomach of each specimen through a ventral
103 vertical incision and subsequently we set all extracted stomach contents into plastic tubes
104 filled with 70% ethanol. Direct extraction of stomach rather than non-lethal techniques (e.g.
105 stomach-flushing) was justified because individuals from our study were also part of research
106 on helminth parasites, the methodology for which requires a complete gastrointestinal
107 analysis and euthanasia.
108 We identified the prey items to the lowest possible taxonomic category following
109 Rafael et al. (2012) and Baccaro et al. (2016). To avoid misclassifications of food items and
110 misestimates in prey sizes we included only intact items for the content analysis (e.g. wings
111 and fragmented exoskeleton parts were not considered in the sample). We measured the Zoology 112 maximum length and width of each prey item with the aidof of millimetric ocular lens to obtain 113 the volume through the Ellipsoid Volume Formula following Griffiths & Mylotte (1987),
114 where l is the prey length and w is the prey width: Journal
115 116 In addition, we calculated the Indexpaper of Relative Importance (IRI), which indicates the 117 importance of each prey category consumed. We calculated it by summing the percentages of
118 number (N%), frequencyNorth-Western (F%), and volume (V%) (as proposed in Biavati et al. 2004): accepted 119
120 To calculate the trophic niche breadth, we used the Levins index (B) described by
121 Pianka (1986), where p is the numerical or volumetric proportion of prey category i and n is
122 the number of prey categories:
123 Use the following type of citation: North-western Journal of Zoology 2021: e211508
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124 The index (B) varies from 1 (specialists in use of food resource) to n (generalists in the
125 use of resources). For better interpretation, we standardized Levin’s measure of niche breadth
126 (Bsta), which limits the value on a scale from 0 to 1 according to the equation below, where n
127 represents the number of resources (prey species) registered. Niche breadth values from 0.0 to
128 0.50 we interpreted as a specialist diet, and values from 0.51 to 1.0 – as a generalist (Hurlbert
129 1978).
130
131 We removed the oocytes from the abdominal cavity of gravid females and preserved
132 them in 5% formalin. We counted the number of oocytes of each female to represent the 133 clutch size. The diameter of 30 oocytes per female were measuredZoology using a stereomicroscope 134 with a millimetric ocular lens. of
135 For the statistical analysis of the results, we performed a Spearman correlation test
136 using BioEstat 5.0 software to determine theJournal relationship between Snout-vent length (SVL)
137 and body mass with volume of the largest prey for each individual. As we were also interested
138 in determining the females’ size and paper the influence of body mass on the clutch size, we 139 performed a Spearman correlation test between female SVL and body mass with number of
140 oocytes and egg diameter.North-Western All variables were tested for normality and homogeneity of 141 variances (significance: accepted p < 0.005). Sexual differences in SVL of males and females were
142 tested using the Student t-test.
143
144 Results
145 We analyzed 100 adult individuals of R. castaneotica, including 77 males and 23
146 females. Males had SVLs ranging from 18.43 to 23.62 mm (mean = 20.27 mm, Standard
147 Deviation = 1.14) and females – from 18.85 to 26.29 mm (mean = 22.69 mm, SD = 1.9).
148 Body mass ranged from 0.3 to 0.9 g (mean = 0.5 g, SD = 0.11) and 0.5 to 2.0 g (mean = 0.85 Use the following type of citation: North-western Journal of Zoology 2021: e211508
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149 g, SD = 0.31) for males and females, respectively. Females were significantly longer (t = -
150 5.5634; p < 0.0001) and heavier (t = -5.0439; p = 0.0001) than males.
151 A total of 34 individuals (34%) had stomach contents, including only two females with
152 prey in their stomach. We identified 31 prey categories belonging to 12 orders (Table 1).
153 Arthropods were the main food resource found in the stomachs of R. castaneotica and the
154 most numerically representative orders were Hymenoptera and Coleoptera. We found 17
155 genera belonging to Formicidae that composed 89.03% of the diet of R. castaneotica. Ants of
156 the genus Crematogaster, Solenopsis and Mayaponera occurred most frequently in the
157 stomachs. The most abundant ant was Crematogaster, while in terms of volume the diet was
158 dominated by larger ants of the genera Cylindromyrmex, Dinoponera and Mayaponera. Plant Zoology 159 material was found in stomachs of females. of 160 The volume of prey consumed did not increase with predator’s SVL and body mass, as
161 we did not find a strong and statistically significant relationship between SVL and prey Journal 162 volume (rs = 0.1306, p = 0.4476) and body mass and prey volume (rs = 0.1557, p = 0.3646).
163 The niche breadth based on prey numbers was 0.24.
164 The number of ovarian eggs rangedpaper from 199 to 720 eggs per female (mean = 403.9,
165 SD = 172.9, total eggs = 3635, n = 9 females with eggs). Egg diameter ranged from 0.73 to
166 1.61 mm (mean = 1.3,North-Western SD = 0.26, n = 270 eggs). We found significant correlation between accepted 167 female mass and egg diameter (rs = 0.7983, p = 0.0099). We did not find significant
168 correlations between female SVL and clutch size (rs = 0.0753, p = 0.8473), body mass and
169 clutch size (rs = 0.4496, p = 0.2246), and egg diameter and SVL (rs = -0.2025, p = 0.6012).
170
171 Discussion
172 We observed sexual size dimorphism in R. castaneotica, in which females are larger
173 than males in body size. Larger females are a common pattern of sexual dimorphism in Use the following type of citation: North-western Journal of Zoology 2021: e211508
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174 anurans, found in almost 90% of species (Shine 1979). Such pattern is often related to sexual
175 selection on fecundity of females, where larger females are favored due to their higher
176 capacity for the allocation and production of larger clutches (Woolbright 1983, Monnet &
177 Cherry 2002, Kupfer 2007). In R. castaneotica, however, the number of oocytes did not
178 increase with female’s body size. Also, the mean clutch size (represented here by number of
179 oocytes) was smaller than those usually reported for bufonids, considering both oocytes and
180 deposited eggs (Crump 1989, Pereyra et al. 2015, Silva et al. 2017). This result agrees with
181 Caldwell & Araujo (2004) that suggested the relatively small clutch size of R. castaneotica
182 from south Amazon is an adaptation to the restricted size of Brazilian nut capsules used as
183 breeding sites. Therefore, we suggest lack of correlation between body size and clutch size in Zoology 184 R. castaneotica in our study is expected due to limitationsof in clutch size and in the number of 185 oocytes produced related to the reproductive mode of the species.
186 On the other hand, we do recognize that the patterns of multiple egg depositions, Journal 187 exhibited by tropical anurans (Prado et al. 2005) display a variation in the number of oocytes
188 produced over the course of a reproductive season (Haddad & Prado 2005, Crump 2015).
189 These variations are particularly expectedpaper in anurans for which reproduction is strongly linked
190 to precipitation, such as bufonids (Caldwell & Araujo 2004, Haddad & Prado 2005). It is
191 likely, therefore, theNorth-Western methodology employed in our study to characterize clutch size (counting accepted 192 oocytes) may have resulted in the lack of, or weak, correlation between female size and egg
193 count, similar to the reasons proposed by Bionda et al. (2011, 2015) for R. arenarum.
194 The narrow niche breadth of R. castaneotica indicates a specialized diet, which is
195 consistent with other studies on Rhinella species that have also documented ants and termites
196 as the dominant prey taxa (Batista et al. 2011, Sabagh et al. 2012, Fajardo-Martínez et al.
197 2013, Flynn et al. 2020). Isacch & Barg (2002) believed that tropical bufonids are ant-
198 specialists, due to their small size and lack of teeth. Furthermore, terrestrial colonial ants, such Use the following type of citation: North-western Journal of Zoology 2021: e211508
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199 as Solenopsis and Blepharidatta, are known to aggregate in enormous colonies that hold large
200 populations above and underneath the soil (Lofgren & Vander Meer 1986, Hölldobler &
201 Wilson 1990) comprising a potential food resource for ground-dwelling anurans. These
202 aspects may explain the high number of these arthropods found in our study.
203 The high incidence of Myrmicinae ants (especially the genus Crematogaster and
204 Solenopsis) have also been reported in the diets of R. icterica, R. crucifer, and R. ornata
205 (Sabagh & Carvalho-e-Silva 2008, Flynn et al. 2020) and for species within R. margaritifera
206 group, such as R. proboscidea and Rhinella sp. (Almeida et al. 2019, Borges et al. 2019). Ants
207 seems to be also a substantial resource in the diet of other ground-dwelling anurans from
208 Eastern Brazilian Amazon, particularly ants of the genus Solenopsis, Pheidolle and Zoology 209 Wasmannia have been commonly reported in the dietsof of anurans from both specialist 210 (Sanches et al. unpublished report, Santana et al. unpublished report) and generalist guilds
211 (Sanches et al. 2019, Corrêa et al. 2020) from Amapá state region. Journal 212 Beetles (Coleoptera) were the second most frequent food item for both males and
213 females. Many studies on Rhinella species have documented the high consumption of beetles,
214 by both sexes (Quiroga et al. 2009, Batistapaper et al. 2011, Maragno & Souza 2011, Sabagh et al.
215 2012). As males spend more energy on vocalization and females need energy for egg
216 production, the differenceNorth-Western between the diets of both sexes reveals the requirements of each accepted 217 sex or different foraging strategies (Maneyro et al. 2004, Brasileiro et al. 2010). In addition,
218 the variation in the diets of both sexes may be caused by different predatory behaviors
219 regarding individual body size, as observed for R. The niche breadth based on prey numbers
220 was 0.24 (as Bufo cf. typhonius in Lima & Magnusson 2000). We also found remains of
221 plants in stomachs, but we did not quantify it. This food item is usually accidentally ingested,
222 as suggested in previous studies on the diet of Rhinella species (Isaacs & Hoyos 2010, Borges
223 et al. 2019). Use the following type of citation: North-western Journal of Zoology 2021: e211508
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224 In the light of our results and based on the diets of other anuran species from this
225 region R. castaneotica is a myrmecophagous consuming the most commonly found genera of
226 ants on the studied area, such as Solenopsis ants. The abundance of arboreal ants, particularly
227 Crematogaster, may be also related to the climbing behavior on understory vegetation already
228 reported for the species (Oliveira-Souza et al. 2020). The small number of oocytes, compared
229 to other bufonid species and the lack of correlation between body size and number of oocytes,
230 may be related to the reproductive mode of the species and needs further investigation to
231 explain its role in sexual size dimorphism of R. castaneotica.
232 Finally, the population of R. castaneotica presented here showed sexual dimorphism
233 (females were larger and heavier than males) and a specialized diet. So far, the present work Zoology 234 is the first to present data on feeding and reproductiveof strategies of R. castaneotica. We 235 highlight the importance of further studies that may evaluate the conservation status and
236 ecological aspects of anurans in the eastern Amazon in Northern Brazil. Journal 237
238 Acknowledgement
239 We thank Universidade Federal do Amapápaper (UNIFAP) and Christopher Jaster (PARNA
240 Montanhas do Tumucumaque) for logistical support during the fieldwork. Specimens were
241 collected under theNorth-Western permission issued by ICMBio (# 48102-2). We thank two anonymous accepted 242 reviewers for suggestions during the reviewing process.
243
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347 Machado, R.A., Bernarde, P.S. (2011): Multiple and heterospecific amplexus between the
348 toads Rhaebo guttatus and Rhinella marina (Anura: Bufonidae). Herpetology Notes 4:
349 167-169.
350 Maneyro, R., Naya, D.E., Da Rosa, I., Canavero, A., Camargo, A. (2004): Diet of the South
351 American frog Leptodactylus ocellatus (Anura, Leptodactylidae) in Uruguay. Iheringia
352 94: 57-61.
353 Maragno, F.P., Souza, F.L. (2011): Diet of Rhinella scitula (Anura, Bufonidae) in the Zoology 354 Cerrado, Brazil: the importance of seasons andof body size. Revista Mexicana de 355 Biodiversidad 82: 879-886.
356 Monnet, J.M., Cherry, M.I. (2002): Sexual size dimorphism in anurans. Proceedings of the Journal 357 Royal Society of London B: Biological Sciences 269: 2301-2307.
358 Oliveira-Souza, A.E., Brito, T.M., Santana, M.M.S., Reis, T.S., Soeiro, A.L.S., Costa-
359 Campos, C.E. (2020): Atelopus paper hoogmoedi (Hoogmoed Harlequin Toad), Rhinella
360 castaneotica (Para Toad), Rhinella lescurei, and Ameerega pulchripecta (Silverstone’s
361 Poison Frog). North-WesternClimbing Behavior. Herpetological Review 51: 813. accepted 362 Peel, M.C., Finlayson, B.L., McMahon, T.A. (2007): Updated world map of the Köppen-
363 Geiger climate classification. Hydrology and Earth System Sciences Discussions 4:
364 439-473.
365 Pereyra, M.O., Candioti, M.F.V., Faivovich, J., Baldo, J.D. (2015): Egg clutch structure of
366 Rhinella rumbolli (Anura: Bufonidae), a toad from the Yungas of Argentina, with a
367 review of the reproductive diversity in Rhinella. Salamandra 51: 161-170. Use the following type of citation: North-western Journal of Zoology 2021: e211508
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368 Pianka, E.R. (1986): Ecology and natural history of desert lizards Lizards – Analysis of the
369 Ecological Niche and Community Structure. 1th Edition. Princeton Legacy Library, New
370 Jersey.
371 Pinto, R.M.C., Menin, M. (2017): Aspects of the natural history of Leptodactylus knudseni
372 Heyer, 1972 (Anura: Leptodactylidae) in a pristine forest in Central Amazonia, Brazil,
373 with comments on ontogenetic variation of its tadpoles. Journal of Natural History 51:
374 2523-2534.
375 Pombal-Junior, J.P., Haddad, C.F.B. (2005): Estratégias e modos reprodutivos de anuros
376 (Amphibia) em uma poça permanente na Serra de Paranapiacaba, Sudeste do Brasil.
377 Papéis Avulsos de Zoologia 45: 201-213. Zoology 378 Prado, C., Toledo, L.F., Zina, J., Haddad C.F.B (2005):of Trophic eggs in the foam nests of 379 Leptodactylus labyrinthicus (Anura, Leptodactylidae): an experimental approach. The
380 Herpetological Journal 15: 279-284. Journal 381 Putnam, R. J. (1994): Community ecology. 1th Edition. Chapman and Hall, London.
382 Quiroga, L.B., Sanabria, E.A., Acosta, J.C. (2009): Size and sex-dependent variation in diet of
383 Rhinella arenarum (Anura: Bufonidae)paper in a Wetland of San Juan, Argentina. Journal of
384 Herpetology 43: 311-317.
385 Rafael, J.A., Melo, G.A.R.,North-Western Carvalho, C.J.B., Casari, S.A., Constantino, R. (2012): Insetos do accepted 386 Brasil: Diversidade e Taxonomia. 1th Edition. Holos, São Paulo.
387 Sabagh, L.T., Carvalho-e-Silva, A.M.P.T. (2008): Feeding overlap in two sympatric species
388 of Rhinella (Anura: Bufonidae) of the Atlantic Rain Forest. Revista Brasileira de
389 Zoologia 25: 247-253.
390 Sabagh, L.T., Carvalho-e-Silva, A.M.P.T., Rocha, C. F. D. (2012): Diet of the toad Rhinella
391 icterica (Anura: Bufonidae) from Atlantic Forest Highlands of southeastern Brazil.
392 Biota Neotropica 12: 258-262. Use the following type of citation: North-western Journal of Zoology 2021: e211508
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393 Sanches, P.R., Santos, F.P., Costa-Campos, C.E. (2019): Diet of the napo tropical bullfrog
394 Adenomera hylaedactyla (Cope, 1868) (Anura: Leptodactylidae) from an urban area in
395 southern Amapá, eastern Amazon. Herpetology Notes 12: 841-845.
396 Shine, R. (1979): Sexual selection and sexual dimorphism in Amphibia. Copeia 1979: 297-
397 306.
398 Silva, M.I., Schiesar, L.C., Menin, M. (2017): The egg clutch and tadpole of Rhinella
399 merianae (Gallardo, 1965) (Anura: Bufonidae) from Central Amazonia, Brazil.
400 Zootaxa, 4294(1): 145-150.
401 Toft, C.A. (1981): Feeding ecology of panamanian litter anurans: patterns in diet and foraging
402 mode. Journal of Herpetology 15: 139-144. Zoology 403 Toledo, L.F., Ribeiro, R.S., Haddad, C.F.B. (2007): Anuransof as prey: an exploratory analysis 404 and size relationships between predators and their prey. Journal of Zoology 271: 170-
405 177. Journal 406 Valentim, D.S.S., Silva e Silva, Y.B., Furtado, M.F.M., Souza, N.S., Costa-Campos, C.E.
407 (2013): Atividade reprodutiva de Rhinella gr. margaritifera (Anura; Bufonidae) em
408 poça temporária no município de paperSerra do Navio, Amapá. Biota Amazônia 3: 188-192.
409 Vargas-Salinas, F. (2007): Breeding Behavior of the Cane Toad Bufo marinus (Bufonidae): A
410 Successfully InvasiveNorth-Western Species. Herpetological Review 38: 12-17. accepted 411 Woolbright, L.L. (1983): Sexual Selection and Size Dimorphism in Anuran Amphibia. The
412 American Naturalist 121: 110-119.
413 Wu, Z., Li, Y., Wang, Y., Adams, M.J. (2005): Diet of introduced bullfrogs (Rana
414 catesbeiana): predation and diet overlap with native frogs on Daishan Island, China.
415 Journal of Herpetology 39: 668-674.
416 Use the following type of citation: North-western Journal of Zoology 2021: e211508
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417 Table 1. Prey categories consumed by R. castaneotica. N = number of individuals; V = total
418 volume of prey; F = frequency of occurrence of each prey category; (%) = percentage; IRI =
419 Index of relative importance.
Males (n = 32) Females (n = 2) Prey item N (%) F (%) V (%) IRI N (%) F (%) V (%) IRI Acari 13 (1.49) 1 (0.67) 1.7 (0.06) 0.74 — — — — Araneae 7 (0.80) 3 (2.00) 1.2 (0.04) 0.95 — — — — Coleoptera (Curculionidae) 24 (2.76) 14 (9.33) 156.09 (5.09) 5.75 4 (100) 2 (100) 79.68 (100) 100 Coleoptera larvae 8 (0.92) 7 (4.67) 18.52 (0.60) 2.07 — — — — Diptera 2 (0.23) 2 (1.33) 5.76 (0.19) 0.59 — — — — Diptera larvae 1 (0.11) 1 (0.67) 1.67 (0.05) 0.28 — — — — Gastropoda 1 (0.11) 1 (0.67) 21.44 (0.7) 0.50 — — — — Hemiptera 2 (0.23) 2 (1.33) 8.09 (0.26) 0.61 — — — — Hymenoptera Zoology (Formicidae) 779 (89.5) 106 (71.1) 2614 (85.3) of81.6 — — — — Dolichoderinae Dorymyrmex sp. 11 (1.26) 3 (2.00) 69.41 (2.26) 1.85 — — — — Dorylinae Journal Cylindromyrmex sp. 26 (2.99) 4 (2.67) 1299.24 (42.3) 16.0 — — — — Ectatomminae Ectatomma 10 (1.15) 7 (4.67) 132.15 (4.31) 3.39 — — — — Myrmicinae paper Acanthognathus 21 (2.41) 3 (2.00) 7.47 (0.24) 1.56 — — — — Acromyrmex 87 (9.99) 8 (5.33) 37.56 (1.22) 5.53 — — — — Blepharidatta North-Western94 (10.7) 7 (4.67) 32.68 (1.07) 5.52 — — — — Crematogaster 247 (28.3)accepted 27 (18) 138.5 (4.52) 17.01 — — — — Lachnomyrmex 20 (2.30) 3 (2.00) 48.87 (1.59) 1.97 — — — — Ochetomyrmex 9 (1.03) 1 (0.67) 2.01 (0.07) 0.59 — — — — Solenopsis 78 (8.96) 17 (11.3) 78.49 (2.56) 7.64 — — — — Sericomyrmex 18 (2.07) 5 (3.33) 37.33 (1.22) 2.21 — — — — Strumigenys 23 (2.64) 3 (2.00) 4.76 (0.16) 1.60 — — — — Pheidole 25 (2.87) 4 (2.67) 53.88 (1.76) 2.44 — — — — Paraponerinae Paraponera 6 (0.69) 1 (0.67) 3.36 (0.11) 0.49 — — — — Ponerinae Centrormyrmex 10 (1.15) 1 (0.67) 48.54 (1.58) 1.13 — — — — Dinoponera 44 (5.05) 4 (2.67) 340.63 (11.1) 6.28 — — — — Use the following type of citation: North-western Journal of Zoology 2021: e211508
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Mayaponera 50 (5.74) 8 (5.33) 280.05 (9.13) 6.75 — — — — Isopoda 14 (1.61) 4 (2.67) 73.46 (2.39) 2.23 — — — — Isoptera 13 (1.49) 2 (1.33) 16.86 (0.55) 1.13 — — — — Neuroptera 1 (0.11) 1 (0.67) 10.05 (0.33) 0.37 — — — — Orthoptera 4 (0.46) 4 (2.67) 136.25 (4.44) 2.53 — — — — Scorpiones 1 (0.11) 1 (0.67) 0.86 (0.03) 0.27 — — — — 420
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North-Western accepted