JHR 97: 505-511 (2024) ore JOURNAL OF 4 ?sreewed oneness oural doi: 10.3897/jhr.97.123968 SHORT COMMUNICATION ) Hymenopter a The International Society of Hymenopterists RESEARCH https://jhr.pensoft.net Gynandromorph records of Melissodes trinodis and Melissodes communis (Hymenoptera, Apidae) from North Dakota, USA Joshua W. Campbell", C. K. Pei? , Karen W. Wright? | USDA-ARS Pest Management Research Unit, Northern Plains Agricultural Research Laboratory, Sidney, MT, 59270, USA 2. University of North Dakota, Earth System Science and Policy, Grand Forks, ND 58201, USA 3 Washington State Department of Agriculture, Olympia, WA 98504-2560, USA Corresponding author: Joshua W. Campbell (joshua.campbell@usda.govy) Academic editor: Jack Neff | Received 26 March 2024 | Accepted 12 May 2024 | Published 6 June 2024 Attps://zoobank. org/707504EB-9077-4B 13-9FE5-9F20523B6357 Citation: Campbell JW, Pei CK, Wright KW (2024) Gynandromorph records of Melissodes trinodis and Melissodes communis (Hymenoptera, Apidae) from North Dakota, USA. Journal of Hymenoptera Research 97: 505-511. https:// doi.org/10.3897/jhr.97.123968 Abstract Bees that express both male and female characteristics are known as gynandromorphs. Here we report and describe two specimens that represent the first documented gynandromorphs of Melissodes trinodis Robertson and one specimen of Melissodes communis Cresson (Hymenoptera: Apidae) that represents only the second known case. All three specimens were collected in North Dakota, USA and exhibit a mosaic pattern of gynandromorphy. Keywords Apoidea, Eucerini, gynandromorph, morphology Introduction Gynandromorphs are individuals that are usually genetic chimeras expressing both male and female characteristics and this phenomenon has been found in most in- sect orders (Pereira et al. 2010; Lightburn et al. 2022). Within bees (Hymenoptera: Apoidea: Anthophila), approximately 140 gynandromorph specimens are described * These authors contributed equally to this work. Copyright: This is an open access article distributed under the terms of the CCO Public Domain Dedication. 506 Joshua W. Campbell et al. / Journal of Hymenoptera Research 97: 505-511 (2024) from all bee families (Lightburn et al. 2022). Although documented in over 30 bee genera, gynandromorphy has been reported more frequently in Andrena Fabricius (Hymenoptera: Andrenidae) and Megachile Latreille (Hymenoptera: Megachilidae) (Michez et al. 2009; Hinojosa-Diaz et al. 2012; Parys et al. 2022). Gynandromorphy has been documented in many different forms in bees but is most commonly found as mosaics where no clear distribution of male and female char- acteristics can be discerned (Wcislo et al. 2004). Other less common gynandromorph patterns documented in bees include male and female characteristics separated bilater- ally, transversally, and various anterior/posterior separations (Dalla Torre and Friese 1899; Wcislo et al. 2004; Michez et al. 2009). Although Melissodes are one of the more commonly collected apids among research- ers in North America, only one known case of gynandromorphy has been documented in the scientific literature (Cockerell 1906). This specimen mentioned by Cockerell (1906), Melissodes hortivagans Cockerell (=/. communis Cresson), was collected in Fedor, TX and described as a ‘partial gynandromorph with a half yellow (right side) and black (left side) clypeus and labrum. Here we report one individual of M. communis and two indi- viduals of M. trinodis Robertson from North Dakota, USA exhibiting gynandromorphy. All three individuals exhibited the mosaic pattern described by Michez et al. (2009) or would be partial bilaterals in the system of Dalla Torre and Friese (1899). Methods The M. communis specimen was collected in a pitfall trap (473 ml Solo cup filled half of the depth with a 50% propylene glycol/water solution) that was active for 7 days (June 10-17, 2021) in the Little Missouri National Grasslands, Billings County, North Da- kota (47.30828, -103.45804). Both M. trinodis specimens were collected with colored (blue, yellow, and white) pan traps (473 ml pan trap filled with soapy water) that were active for 24 hours. The M. trinodis specimens were part of a greater survey effort to sample grassland bee communities in North Dakota (Pei et al. 2022). One gynandro- morph was collected in a sampling event on July 26, 2017 from Long Lake National Wildlife Refuge, Burleigh County, North Dakota (48.81845, -100.22222). A second gynandromorph was collected on a July 15, 2018 sampling event in Agnes Marsh Wa- terfowl Production Area, Grand Forks County, North Dakota (48.08277, -97.72434). Here, we describe each specimen and provide detailed photographs of the external morphology. For each specimen, the following external morphological measurements were made: total body length (mm), head width (taken from widest part of eyes) (mm), intertegular distance (ITD) (mm), abdominal width (taken from widest part of abdo- men) (mm), scape + pedicel length (mm), and flagellum length (mm) (Table 1). All photographs of M. trinodis were taken with a Canon EOS 6D Mark II Digital SLR and photographs of M. communis with a Leica DMC 4500. The M. communis specimen is housed at the USDA Northern Plains Agricultural Research Laboratory native bee collection (Sidney, MT) and the M. trinodis specimens are located at North Dakota State University in the School of Natural Resources Sciences (Fargo, ND). Gynandromorphs of Melissodes communis and M. trinodis 507 Table |. Morphological measurements (mm) of total body length, head width, ITD, abdomen width, scape + pedicel length, and flagellum length of M. communis and M. trinodis. GF= Grand Forks specimen, BC= Burleigh County specimen. Melissodes communis Melissodes trinodis (GF) Melissodes trinodis (BC) Total Body Length 11.26 11.49 P21 Head Width 3.81 Duro 3.86 HED 2.4 2sFi, oa! Abdomen Width 4.01 4.3 4.5 Scape + Pedicel Length 0.78 0.98 0.95 Flagellum Length 2:91 N/A Qe Results Melissodes communis This specimen was collected along with 106 other Melissodes specimens collected in the same 2021 sampling effort. The specimen exhibits primarily bilateral asymmetric yellow male coloration (right side) and dark female (left side) coloration of clypeus and labrum (Fig. 1A; labrum not visible in photo). Both antennae have 12 segments. Other than the clypeus, labrum and antennae, specimen appears to be male (e.g., legs lacking tibia scopal hair, exhibiting male genitalia) (Fig. 1B—D). Melissodes trinodis These specimens were collected along with 1,252 (Burleigh specimen) and 1,126 (Grand Forks specimen) Melissodes specimens in the same year sampling effort. Both specimens exhibit bilateral asymmetric male/female (yellow/dark) coloration of cl- ypeus and labrum, with male-associated yellow coloration found on the left side for the Grand Forks specimen and on the right side for the Burleigh specimen (Figs 2A, 3A). All other external characteristics of each specimen appear female (Figs 2B—D, 3B—D). However, the antennae were not present for the specimen from Burleigh County so we cannot ascertain the number of antennal segments. Additionally, one hind tibia is missing from the Grand Forks specimen and, thus, we cannot determine if scopal hairs were present. Discussion We describe the first gynandromorphs from Melissodes trinodis and the second of M. communis. Other than Cockerell (1906), these are the first reports of gynan- dromorphy from this genus in the scientific literature, and all three specimens appear to follow the mosaic pattern described by Michez et al. (2009). However, M. communis and one specimen of M. trinodis (specimen from Grand Forks Coun- ty) display a ‘patchiness’ of the yellow coloration on their clypeus (see Figs 1A, 2A). This patchiness is also seen on the labrum for M. trinodis. Due to the lack 508 Joshua W. Campbell et al. / Journal of Hymenoptera Research 97: 505-511 (2024) - WY (i) 7a BS ATH \ —| 5 ge ey Gah ey : > Ter Oe Figure |. Gynandromorph of Melissodes communis collected from Billings County, North Dakota, USA A face B dorsal view C lateral view, and D male genitalia (partially exposed). of sharp morphological distinctions of male and female characters (e.g., clypeus yellow color not uniform), these two specimens could be considered mosaics by Michez et al. (2010) or would be called partial bilaterals in the system of Dalla Torre and Friese (1899). Both of the bee species in this study are widely distributed in North America, with M. communis found throughout the United States, southern Canada and Mexico, whereas WM. trinodis is primarily found in the eastern half of the United States but can also be found in Canada and Mexico. Melissodes communis is polylectic, feeding on mul- tiple pollen sources, whereas M. trinodis is primarily considered an Asteraceae specialist and one of the main visitors of flowering cultivated sunflower (Portlas et al. 2018). Jones et al. (2021) was able to observe a gynandromorph bee (Xenoglossa pruinosa Say (Hymenoptera: Apidae)) foraging prior to collection, and, thus, was able to glean some behavioral information. Since the specimens described here were collected in passive traps, no behavioral observations could be collected. Prior to these specimens, only eight other gynandromorphic individuals from tribe Eucerini had been documented, (Cockerell 1906; Jones et al. 2021; Parys et al. 2022), suggesting gynandromorphs may be rare within Eucerini compared to Gynandromorphs of Melissodes communis and M. trinodis 509 1.00 mm el 1.00 mm re Figure 2. Gynandromorph of Melissodes trinodis collected from Grand Forks County, North Dakota, USA A face B dorsal view C€ lateral view, and D abdomen. other bee groups. Alternatively, there could be a bias in finding/collecting gynandro- morphs from Eucerini or there is a lack of reporting them. In addition to Melissodes and Xenoglossa, other Eucerini genera in which gynandromorphs have been docu- mented are Alloscirtetica Holmberg (Hymenoptera: Apidae) (Urban 1999), Flori- legus Robertson (Hymenoptera: Apidae) (Parys et al. 2022), and Tetralonia Spinola (Hymenoptera: Apidae) (Dalla Torre and Friese 1899). It is unclear why gynandro- morphy is uncommon or poorly documented within Eucerini. However, numerous explanations are thought to cause gynandromorphy in bees (or other arthropods) including various developmental mechanisms (Jones et al. 2021), endosymbiotic bacteria (Narita et al. 2010), epigenic causes (Sommaggio et al. 2021), and nu- merous environmental stressors such as temperature (Drescher and Rothenbuhler 1963) and pollutants (Dantchenko et al. 1995; Olmstead and LeBlanc 2007). The presence of gynandromorphs in wild bee populations is considered rare and the causes unknown and, thus, further research and documentation is widely needed to elucidate the causes of gynandromorphy (Jones et al. 2021; Parys et al. 2022). There has been an increase of documentation of gynandromorphy in wild bees in recent literature (Jones et al. 2021), but whether this is an upsurge of interest by the sci- entific community to document this occurrence or an increase in a causation of this phenomenon remains unknown. 510 Joshua W. Campbell et al. / Journal of Hymenoptera Research 97: 505-511 (2024) 1.00 mm mi 1.00 mm rm Figure 3. Gynandromorph of Melissodes trinodis collected from Burleigh County, North Dakota, USA A face B dorsal view C lateral view, and D abdomen. Acknowledgments We thank Fraser Watson and Niall Horton for pitfall collection and Cody Meservy for providing photographs and morphological measurements and Alex Morphew for speci- men processing. 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