Short Communication Journal of Orthoptera Research 2021, 30(1): 27-30 Description and photographs of cricket parental care in the wild DARIN J. MCNEIL! 24, BETTINA ERREGGER?* 1 Department of Natural Resources, Cornell University, Ithaca, NY 14850, USA. 2 Cornell Laboratory of Ornithology, Ithaca, NY 14850, USA. 3 Institute of Biology, University of Graz, Graz, Austria. 4 Current address: Pennsylvania State University, Department of Entomology, University Park, PA 16802, USA. 5 Current address: Institute of Animal Nutrition, Products and Nutritional Physiology, University of Natural Resources and Life Science, Vienna, Austria. Corresponding author: Darin J. McNeil (darin.j.mcneil@gmail.com) Academic editor: Kevin Judge | Received 14 March 2020 | Accepted 12 June 2020 | Published 23 March 2021 http://zoobank.org/C99D80AF-2002-464F-B990-17BCIED30C32 Citation: McNeil DJ, Erregger B (2021) Description and photographs of cricket parental care in the wild. Journal of Orthoptera Research 30(1): 27-30. https://doi.org/10.3897/jor.30.52079 Abstract Although certain forms of parental care are relatively widespread phe- nomena among insects, within Orthoptera, parental care is rare. Short- tailed burrowing crickets (Anurogryllus spp.) are among the few members of this order for which extensive parental care has been documented. However, accounts of parental care in Anurogryllus have been largely under laboratory conditions, and observations of this behavior in the wild are rare. Herein we present photographic observations from a mountain slope in Honduras where we discovered an active Anurogryllus brood chamber where an adult female was tending her brood. We present these results in the context of parental care in insects and compare our observations with those reported in past literature published on Anurogryllus crickets’ parental behavior. Keywords behavior, eggs, Honduras, nymphs, Orthoptera, short-tailed cricket Introduction Parental care, though widespread and important across many vertebrate taxa (Balshine 2012), is broadly rare among inverte- brates (Irumbo 2012). In insects, parental care has evolved in- dependently several times (e.g., in Hymenoptera, Coleoptera, Blattodea, Dermaptera) but is thought to be largely absent from other insect orders. In Orthoptera, parental investment takes one of two main forms: pre-mating and post-mating (Gwynne 1983). Pre-mating parental investment is common in Orthoptera (espe- cially Ensifera; Gwynne 1983) whereby males provide their mates with a nutrient-rich spermatophylax, which is an appendage of spermatophores often containing a high proportion of protein. Post-mating parental investment (i.e., parental care), on the other hand, is much less common in Orthoptera and varies from dig- ging burrows to protect nymphs from predators (e.g., Gryllotal- pidae, Phaneropterinae: Gwynne 1983) to the protection of eggs (e.g., Neotridactylus apicialis (Say, 1825): Gwynne 1983) to mater- nal care for their young (e.g., Anurogryllus muticus (De Geer, 1773): West and Alexander 1963, Lee and Loher 1995; Hemiandrus spp.: Gwynne 2004). Despite being uncommon in Orthoptera, post- mating parental care occurs more commonly in a variety of re- lated insect orders as a means to maximize juvenile survival (e.g., Blattodea: Nalepa 1988; Dermaptera: Kolliker 2007). For example, the wood roach Cryptocercus punctulatus Scudder, 1862 provides bi- parental care to nymphs for as long as three years (Nalepa 1988). Among Orthoptera, crickets in the genus Anurogryllus Saussure, 1877 provide exceptional parental care to their young, with care frequently lasting for several weeks (West and Alexander 1963, Weaver and Sommers 1969). As first detailed by West and Alexan- der (1963) and Weaver and Sommers (1969), Anurogryllus muticus females extensively clean their eggs, provision nymphs with forage collected outside the burrow and trophic eggs, and vigorously de- fend young from males and other intruders to their burrow (West and Alexander 1963). Although parental care in Anurogryllus spp. is well described in captivity, few accounts of parental care in this unique orthopteran genus exist from the wild, and no published photographs exist of natural Anurogryllus brood chambers where this rare example of orthopteran parental care is known to occur. This is an important gap in the literature as species behaviors in the wild may differ drastically from those observed in captivity (Alvarez and Nicieza 2003, Tenger-Trolander et al. 2019). For the first time herein, we provide photographs of the natural brood chamber of an Anurogryllus cricket observed in a mountain rain forest in northern Honduras. Materials and methods Our observations were made on 1 February 2017 at Rio Santiago Nature Reserve in San Francisco, Atlantida, Honduras (15.592223°N, -87.062514°E; 150 m) at 20:55 (Fig. 1). This re- gion is heavily forested and dominated by contiguous broadleaf JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) 28 evergreen forest. On the evening when our observations were made, the temperature was approximately 19°C and the sky was overcast with light rain. Our observation occurred during a rep- tile/amphibian survey through the nature reserve during which we rolled stones and other objects to document resident wildlife. During this survey, we rolled a small stone with dimensions ap- proximately 7.5 x 13 cm. This stone, which was sitting atop the soil within the forest path, was covering the brood chamber of an Anurogryllus cricket (Fig. 1). Female Anurogryllus are readily identified by their habitus in general (e.g., wing structure), but es- pecially by their greatly reduced ovipositor, consistent with their common name: the short-tailed cricket (Weaver and Sommers 1969). They are also the only known Gryllids that exhibit parental care of nymphs (West and Alexander 1963, Weaver and Sommers 1969), making brooding females very easy to identify. With that in mind, and because detailed descriptions of Anurogryllus species present in Honduras are lacking, we did not attempt to identify the crickets observed at species level. Four photographs were taken to document the observation. The record was also posted to the citizen science platform iNaturalist (https://www.inaturalist.org/ observations/5089680). Results The main brood chamber was ~57 mm long by ~23 mm wide (Fig. 2) and included a single entrance (~31 mm in length) fac- ing west that allowed travel between the burrow and the soil surface. This brood chamber contained 1) an adult female An- urogryllus cricket, 2) three Anurogryllus nymphs (first instar), and 3) three Anurogryllus eggs. We did not observe Anurogryllus nymphs attached to the underside of the rock and thus believe this was the complete brood (three nymphs, three eggs). During our brief encounter with these crickets, including photograph- ing them from several angles, the female was reluctant to expose DJ. MCNEIL AND B. ERREGGER the nymphs and made no attempt to abandon the brood. After documenting the event, the stone was carefully replaced, and the crickets were not harmed. Discussion Parental care is common in some insect orders (e.g., Hyme- noptera) but remarkably rare in others (e.g., Orthoptera: Trumbo 2012). Among the best documented cases of extensive orthopteran maternal care are the ground weta (Hemiandrus spp.) and crickets in the genus Anurogryllus (West and Alexander 1963, Smith et al. 2013). We believe our observations constitute the first published photographs of an opened, active Anurogryllus cricket brood cham- ber in the wild. Although Anurogryllus crickets are known to be- come highly abundant in some areas (West and Alexander 1963), nearly all published studies of their breeding behavior have been restricted to captive conditions. This is surprising given that the natural history and ecology of Anurogryllus crickets has been stud- ied for over a half century and the species is widespread across many states and countries (West and Alexander 1963, Erregger and Schmidt 2018). Although we expect broad reproductive behaviors to remain largely consistent between captivity and the wild, insect behavior can be impacted by captive conditions in profound ways (Tenger-Trolander et al. 2019), underscoring the importance of such natural history observations. In many regards, the Anurogryllus sp. brood chamber observa- tion was similar to those reported in the literature for A. muticus. For example, West and Alexander (1963) observed female A. mu- ticus to construct a brood chamber several centimeters deep into loose soil, linked to the surface by a single connecting channel, though these observations all occurred within a captive environ- ment where burrowing depth was constrained. In contrast, Weav- er and Sommers (1969) created plastic casts of A. muticus burrows in the wild and found them to be multi-chambered and as deep Honduras S; alvador Fig. 1. A. Female Anurogryllus cricket with eggs and nymphs observed in northern Honduras; B. Map of northern Honduras; star repre- sents observation location. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) DJ. MCNEIL AND B. ERREGGER 10 20mm Fig. 2. A. Photo of a female Anurogryllus cricket in a brood chamber beneath a stone on a forest path in Honduras; B. Drawing sim- plifying the structure of the brood chamber (scale bar approximated from cricket body size. The brood chamber, shown in light gray, was beneath the footprint of a small stone that was largely in direct contact with the soil (dark gray). The chamber contained an adult female Anurogryllus cricket as well as three eggs (open circles) and three first-instar nymphs (asterisks). Map data 2019 (c) Google. as 50 cm into the soil. In this way, the burrow we observed at Rio Santiago was more similar to those reported by West and Alex- ander (1963) from the lab than those by Weaver and Sommers (1969) in Louisiana, US. Although we did not measure the bur- row’s depth at Rio Santiago, we can safely conclude that it was <5 cm deep and contained only an “upper chamber” (sensu Weaver and Sommers 1969) with no lower chambers (Fig. 2). Our obser- vation that eggs and nymphs co-occuring is also consistent with past literature suggesting that adults lay eggs successionally (i.e., each individually over the course of several days), sometimes even with nymphs in their burrows (Lee and Loher 1995, Erreg- ger personal observation). The Anurogryllus from Honduras was unique from those pre- viously reported for Anurogryllus in several ways: the Rio Santiago brood chamber contained far fewer nymphs (n = 3) and eggs (n = 3; Fig. 2) than those reported by others (up to 129 eggs or 89 nymphs/burrow; Weaver and Sommers 1969). One possible expla- nation for fewer nymphs and eggs (as compared to previous reports by e.g., Weaver and Sommers 1969) might be the habitat context of the brood chamber. Anurogryllus cricket burrows are best known from grassland habitats (Erregger, personal observation), whereas the burrow described here was deep within a closed-canopy for- est. Although Anurogryllus crickets are foraging generalists, ‘typical’ food sources (e.g., seeds, flowers, leaves) may have been limited within the forest, thus limiting reproductive output. This observa- tion is potentially supported by our idea of no vegetative provi- sions within the burrow (Fig. 2). Indeed, A. muticus are known to conspicuously provision their young nymphs with vegetation gath- ered outside the burrow when food is abundant. Vegetational struc- tures are also used to insulate the chamber from intruders during the day and when the female leaves the burrow (Erregger, personal observation from Panama). Thus, other factors, such as predation pressure, also have an impact on their survival rate (Erregger and Schmidt 2018). Furthermore, the geographic location (i.e., eleva- tion level) and the soil characteristics (e.g., for digging) might be re- stricting factors, reducing survival success. The differences we report here for an Anurogryllus brood chamber in the wild as compared to those studied in captivity highlight the importance of wild obser- vation, as natural conditions and ecological context clearly drive at least some variation in life history for short-tailed crickets. Our observations of an Anurogryllus brood chamber provide interesting insights into the parental care behaviors present with- in Orthoptera. Although some aspects of Anurogryllus breeding behavior were confirmed in the wild by our record, others (e.g., brood size, provisioning behavior, etc.) appear to vary among re- gions, species, and/or study context (e.g., lab vs. field). Given this, several important limitations of our observation should be kept in mind. First, the brood chamber we observed represents a sin- gle observation; how typical any components of this observation are in the broader context of the life history of Anurogryllus would require a much more thorough examination. Moreover, the spe- cies identity of the Anurogryllus brood we observed here remains unknown—species-specific life history differences may explain some of the differences we report here as compared to laboratory observations. Still, the different life history pattern we observed in Honduras in contrast to those observed in the lab highlights the need to assess species behavior across multiple contexts, including both in a lab setting and the wild. Acknowledgements We are grateful to L. Symes (Cornell Lab of Ornithology) for helpful edits on early drafts of this manuscript. We appreciate the field support in Honduras from M. L. Kazour (Utica College) and JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(1) 30 R. E. Bennett (Cornell University). 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