Cyclocephala

Cyclocephala is a genus of scarab beetles in the family Scarabaeidae, subfamily Dynastinae, and tribe Cyclocephalini, first described by Dejean in 1821.^[1] Comprising approximately 350 species, it represents one of the largest genera in the subfamily, with ongoing discoveries adding to its diversity.^[2] Commonly known as masked chafers, these beetles are characterized by a robust body, with adults typically measuring 10–25 mm in length, a clypeus that converges to a rounded or emarginate apex, and antennae consisting of 8–10 segments forming a lamellate club.^[1][3] The genus exhibits a broad distribution across the Americas, ranging from southeastern Canada through the United States and the West Indies to Argentina, though the majority of species are concentrated in the Neotropical region, including high diversity in Brazil's biomes such as the Amazon and Atlantic Forest.^[1][2] Some species have been introduced outside their native range, establishing populations in places like Australia and Hawaii.^[4] Ecologically, Cyclocephala species are phytophagous, with larvae (known as white grubs) feeding on roots of grasses and crops, contributing to nutrient cycling in soils.^[2][1] Adults are predominantly nocturnal, often attracted to lights, and Cyclocephala species, along with other cyclocephalines, play a crucial role as pollinators for approximately 900 Neotropical plant species, including those in the families Annonaceae and Araceae, where they exploit thermogenic flowers for mating and feeding.^[1][4] Economically, certain species are significant pests in agriculture and turfgrass, damaging crops like corn and sugarcane through larval root-feeding, prompting management strategies in affected regions.^[3][4]

Description

Adults

Adult Cyclocephala beetles, belonging to the subfamily Dynastinae within the family Scarabaeidae, are medium-sized scarabs characterized by a robust body with a clypeus whose sides converge toward the apex, 8 to 10 antennomeres forming a lamellate club, and a maxilla featuring distinct teeth.^[5] Males exhibit sexual dimorphism, including an enlarged protarsus and a larger median tarsal claw on the forelegs compared to females.^[5] For example, in C. labidion, males measure 14.0–15.5 mm in length and 6.5–7.0 mm in width, with a dark testaceous coloration accented by black on the frons and piceous markings on the pronotum and elytra, while females are slightly smaller at 11.5–13.8 mm long and 5.9–7.6 mm wide.^[5]

Immatures

The immature stages of Cyclocephala beetles encompass eggs, three larval instars, and pupae. Eggs are typically oval or elliptical, measuring 1.0–1.5 mm in length and 0.8–1.2 mm in width at oviposition, with a milky white chorion that darkens to brownish-yellow near hatching. A notable feature in many species is significant embryonic expansion, where eggs increase in volume up to threefold due to water uptake.^[6][7][8] Larvae are scarabaeiform, C-shaped when at rest, and creamy white with a darkened, prognathous head capsule; they possess three instars, determined by head capsule width following Dyar's rule (approximately 1.6-fold increase per instar). The first instar is small (head width ~1.1–1.6 mm) and translucent. The second instar (head width ~1.8–2.6 mm) has increased setation and mobility. The third instar (head width ~2.8–4.0 mm, body length up to 30–40 mm) is the largest. Morphological diagnostics include a raster pattern on the terminal abdominal segment featuring bifurcate or halberd-shaped setae, varying by species—for instance, C. paraguayensis third instars have distinctly bifurcated raster setae, while C. melanocephala exhibit 3–4 dorsoepicranial setae per side and palidia with 3–4 bifurcate setae.^[9][6][10] Prior to pupation, third-instar larvae form a prepupal stage, during which they construct earthen cells 10–20 cm deep in the soil. Pupae are exarate, elongate-oval, and glabrous or lightly pubescent, with lengths of 12–20 mm; the integument is initially whitish, turning yellowish. Key features include dioneiform organs (paired sensory structures) on abdominal tergites I–VI, numbering two to five per intersegmental membrane, aiding in gas exchange. Species-specific traits, such as the smooth integument and short pubescence in C. melanocephala pupae (15.2–15.4 mm long), facilitate identification within the genus.^[9][10][7]

Taxonomy

Etymology and History

The genus Cyclocephala was established by French entomologist Pierre François Marie Auguste Dejean in 1821 within the first edition of his Catalogue de la collection des coléoptères de M. le Baron Dejean. Dejean's publication included species such as Melolontha geminata Fabricius and Scarabaeus barbatus Fabricius, which rendered the genus-group name available from that date despite initial nomenclatural uncertainties.^[4] Early taxonomic work on cyclocephaline scarabs traces to Carl Linnaeus, who described Scarabaeus amazonus in 1767—the species later designated as the type for Cyclocephala—and Johan Christian Fabricius, who in 1801 classified 11 cyclocephaline species across several genera, including what would become Cyclocephala. Confusion arose regarding authorship, with some attributing the genus to Pierre André Latreille in 1829, but modern consensus confirms Dejean's priority, as clarified by Bousquet and Bouchard (2013). The original type specimen of S. amazonus is lost, prompting Sebő Endrődi to designate a neotype from Suriname in 1966.^[4] The tribe Cyclocephalini, encompassing Cyclocephala and 13 other genera, has a complex history reflecting broader classifications within Dynastinae. Early groupings appeared in works by Hermann Burmeister (1847, 1855), who treated cyclocephalines as a distinct section of Melolonthidae. The tribe name was formally proposed in its modern sense by William Sharp MacLeay (1819) for Australian forms, but Thomas Say (1835) and Francis Polkinghorne Hope (1841) applied it to New World dynastines, solidifying its usage. Endrődi's comprehensive monograph on Dynastinae (1966) recognized 14 genera in the tribe, emphasizing its predominantly Neotropical distribution and primitive traits relative to other dynastines; subsequent revisions, such as those by Ratcliffe (2003) for Central America and Ratcliffe and Cave (2006) for the broader group, have expanded species counts and refined boundaries.^[4][11]

Species Diversity and Groups

The genus Cyclocephala is one of the most species-rich taxa within the subfamily Dynastinae, comprising approximately 340 described species. This diversity is predominantly concentrated in the Neotropical region, where the genus exhibits its highest endemism and abundance, with 81 species recorded in Peru alone.^[12] Within Peru, the departments of Cusco (42 species), Madre de Dios (31 species), and Pasco (27 species) represent key centers of diversity, reflecting the genus's adaptation to varied tropical ecosystems.^[12] A smaller number of species extend into the Nearctic region, including several in North America, while the genus is absent from the Palearctic and Oriental realms.^[4] Taxonomic organization within Cyclocephala has historically relied on morphological characters, particularly those of the head, to delineate species groups. Early classifications by Burmeister (1847) divided the genus into eight informal species groups, a framework later refined by Endrődi (1966) based on detailed examinations of adult morphology.^[4] These groups are:

• Cyclocephalae anomalinae
• Cyclocephalae acutae
• Cyclocephalae parabolicae
• Cyclocephalae heterocerae
• Cyclocephalae reflexae
• Cyclocephalae microcephalae
• Cyclocephalae sinuatae
• Cyclocephalae eurycephalae

This grouping emphasizes variations in head shape, frons structure, and clypeal margins, aiding in species identification amid the genus's cryptic diversity.^[4] Subsequent revisions, such as those focusing on regional faunas (e.g., black species in Colombia), have proposed additional subgroupings based on coloration, genitalia, and elytral patterns, but the Burmeister-Endrődi system remains foundational for broader taxonomic work.^[13] Ongoing molecular studies continue to refine these boundaries, revealing potential cryptic species within established groups.^[4]

Distribution and Habitat

Geographic Range

The genus Cyclocephala is endemic to the Americas, with a distribution spanning the Nearctic and Neotropical realms. It occurs from southeastern Canada southward to Argentina, encompassing a wide latitudinal range across North, Central, and South America, as well as the West Indies.^[12] This broad range reflects the genus's adaptability to diverse environments, though most species are concentrated in tropical and subtropical zones.^[11] In the Nearctic region, Cyclocephala species are primarily found in the southern and eastern United States, with records extending into Mexico and parts of Canada. For instance, species like C. lurida are documented in north-central U.S. states such as Wisconsin and Nebraska.^[14] The genus reaches its highest diversity in the Neotropical realm, particularly in northern South America, with over 350 species described for the genus overall. Countries with significant species richness include Brazil, Colombia, Venezuela, Peru, and Ecuador, often associated with forested and agricultural habitats.^[15] Central America serves as a transitional zone, with numerous species recorded in nations like Panama, Costa Rica, Guatemala, and Honduras, contributing to the genus's overall connectivity across the continents. In the Caribbean, occurrences are noted in the Bahamas, Dominican Republic, and Puerto Rico, though less diverse than mainland populations.^[4] This pan-American distribution underscores Cyclocephala's role in Neotropical ecosystems, with ongoing discoveries expanding known ranges in understudied areas.^[16]

Habitat Preferences

Cyclocephala species exhibit diverse habitat preferences across tropical, subtropical, and temperate regions, primarily in the Americas, with some introduced elsewhere. The genus is most abundant in Neotropical biomes such as the Amazon rainforest, Atlantic Forest, and Cerrado savanna, where species richness is highest, with high endemism, for example over 70% of species collected across Brazilian biomes being endemic to the Amazon. In these environments, adults are often associated with forested or fragmented woodland areas, while larvae inhabit soils beneath vegetation. Outside the Neotropics, North American species favor temperate and transitional zones, including agricultural fields and urban landscapes.^[2][17] Larval stages predominantly occupy underground habitats in well-drained soils rich in organic matter, typically at depths of 5–25 cm, where they feed on roots of grasses, crops, and decaying plant material. Preferred soil conditions include moist environments with adequate aeration, as eggs and early instars are sensitive to desiccation and high temperatures above 40°C or moisture below 8%. In turfgrass systems, such as those dominated by bermudagrass or cool-season varieties, larvae thrive in the root zone, contributing to their role as pests in lawns and golf courses across the southern and midwestern United States. In agricultural settings, they are common under corn, sorghum, sugarcane, and pasture grasses, with densities reaching up to 160 larvae per square meter in Uruguayan pastures. In tropical rainforests, larvae develop in litter layers or near host plant roots, such as those of Annonaceae or palms.^[18][6][19] Adult Cyclocephala beetles are nocturnal and diurnal refuges in soil or leaf litter, emerging to forage in open or canopy habitats linked to floral resources. In the Atlantic Forest of Brazil, species like Cyclocephala distincta are frequently encountered in lowland dense rainforests, visiting Neotropical palms (Arecaceae) for pollination and feeding. In Venezuelan lowland tropical rainforests, adults exploit canopy flowers and extrafloral nectaries of trees such as Albizia pedicellaris and Matayba guianensis, indicating a preference for diverse angiosperm communities. Temperate species, such as Cyclocephala lurida in the southern U.S., inhabit grasslands, turfs, and disturbed areas, while some adapt to non-forested biomes like the Caatinga or Pampa in Brazil, though with lower abundances compared to forested zones. Overall, habitat selection reflects a balance between larval soil requirements and adult associations with thermogenic or odor-emitting plants.^[6][20][17]

Life History

Life Cycle Stages

Cyclocephala beetles exhibit a complete metamorphosis, progressing through four distinct life stages: egg, larva, pupa, and adult. Most species in the genus are univoltine, completing their life cycle in approximately one year, though durations vary by species, temperature, and geographic location. For instance, the life cycle of Cyclocephala distincta averages 108 days under laboratory conditions, while C. celata requires about 164 days in captivity.^[6][21] Adults are the reproductive stage, emerging from the soil in mid- to late spring or early summer, typically from late May to mid-July depending on latitude. In central Texas, peak adult flights occur in mid- to late June, with males emerging first and becoming active at dusk, attracted to lights. Females follow shortly after, releasing pheromones to attract mates; mating occurs at night on or near the soil surface. Adults do not feed significantly and live for 2–3 weeks, during which females oviposit 11–15 eggs on average, burying them 2–6 inches deep in moist soil near grass roots. Species like C. borealis and C. lurida in Ohio lay eggs in July, while C. distincta females produce fertile eggs with an 81% fertility rate over a 3–4 day period.^[19][22][6] The egg stage lasts 10–18 days, during which the pearly white, oval eggs (initially 1–1.2 mm long) absorb moisture and swell to about 2 mm in diameter. Hatching success is high, reaching 74–93% in studied species; for C. distincta, eggs hatch in 11 days at around 25°C, while C. celata eggs take 14 days and are laid in spherical soil chambers. Upon hatching, neonate larvae are small, C-shaped grubs measuring 4–5 mm long.^[22][6][21] Larvae, known as white grubs, undergo three instars and are the primary feeding and damaging stage, consuming roots of grasses and other plants. First-instar larvae (head capsule ~1.4–1.6 mm wide) last 18–25 days and feed near the soil surface in summer; second instars (~2.6 mm head capsule) persist for 17–26 days with similar habits. The third instar, the largest at 20–25 mm long, dominates the cycle, lasting 40–68 days initially before overwintering; grubs burrow 3–12 inches deep in fall, hibernate through winter, and resume feeding in spring. In C. borealis and C. lurida, third-instar damage peaks in September, with survival rates across instars ranging from 22–97% in C. distincta. Head capsule widths follow Dyar's rule, increasing by about 1.6 times per instar.^{[22][6][21][19]} Pupation occurs in late spring or early summer, 3–6 inches below the soil surface in earthen cells formed by the mature third-instar larva. The pupal stage, a non-feeding transformative phase, lasts 11–38 days; pupae are initially creamy white, turning reddish-brown, and measure about 17 mm long by 8 mm wide in northern species. For C. distincta, pupation takes 11 days following a 4-day pre-pupal period, with a 67% emergence rate, while C. celata pupae develop over 38 days in enclosed cells. Successful pupae eclose into adults, which remain in the pupal chamber until dusk before emerging to initiate the next generation.^{[22][19][6][21]}

Development and Reproduction

Cyclocephala species undergo complete metamorphosis, featuring a holometabolous life cycle with distinct egg, larval (three instars), pupal, and adult stages. The genus typically completes its development in one year, distinguishing it from related genera like Phyllophaga that often require two to three years. Adults emerge from pupal cells in the soil during late spring or early summer, depending on the species and region, and live for several weeks to facilitate reproduction. Larvae, known as white grubs, are the primary feeding stage, consuming roots and organic matter underground, while adults rarely feed or do so minimally on nectar or pollen.^[23][24] Developmental durations vary by species and environmental conditions, but representative data from Cyclocephala barrerai illustrate the process: eggs hatch in approximately 22 days, first-instar larvae last 36 days, second-instar 37 days, third-instar 93 days, prepupal stage 36 days, and pupal stage 20 days, totaling about 277 days under controlled conditions. In Cyclocephala celata, the full cycle from egg to adult spans 164 days in captivity, with eggs incubating for 14 days, larval instars totaling around 117 days, and pupation lasting 38 days. Across the genus, egg-to-adult development ranges from 108 to 269 days for eight studied species, influenced by soil temperature and moisture. Third-instar larvae construct earthen cells for pupation, where they transform into adults that remain until emergence.^[24][25][24] Reproduction in Cyclocephala is sexual, with adults mating shortly after emergence, often nocturnally and guided by female-released sex pheromones produced by symbiotic bacteria or cuticular hydrocarbons. In C. barrerai, mating progresses through premating (male searching and antennating), mating (mounting and aedeagus insertion with antennal stimulation), and postmating (male guarding, lasting longer in virgin males at about 388 seconds). Sexual activity begins around 25 days post-emergence, with males active every 2–3 days and females every 4–8 days; mated females oviposit fertilized eggs in soil chambers. Females typically lay 20–31 eggs over their lifespan, as seen in C. barrerai, with high egg viability (up to 93% in C. celata). Most species are univoltine, producing one generation per year, though C. lunulata is bivoltine. Sex ratios are generally 1:1 but can skew toward females in later generations under rearing conditions.^{[24][26][26][24][25][24]}

Ecology

Diet and Feeding

Cyclocephala larvae, commonly known as white grubs, are primarily root feeders that inhabit the soil, where they consume the roots of grasses and other plants, as well as decaying organic matter and thatch. This subterranean feeding occurs across multiple instars, with the third instar causing the most significant damage due to increased size and voracity, particularly in late summer and fall when larvae are actively growing. Preferred hosts include turfgrasses such as Kentucky bluegrass (Poa pratensis), tall fescue (Festuca arundinacea), and bermudagrass (Cynodon dactylon), as well as field crops like corn (Zea mays), wheat (Triticum aestivum), and oats (Avena sativa).^[22][27][18] In rearing conditions, larvae of species like Cyclocephala barrerai thrive on a substrate enriched with pulverized wood, dry leaves, and grass roots such as Brachiaria decumbens, exhibiting rhizophagous and saprophytic habits that allow them to digest both living roots and decomposing plant material.^[24] Adult Cyclocephala beetles exhibit nocturnal feeding behaviors, primarily visiting flowers to consume pollen, nectar, and occasionally floral tissues, which supports their energy needs for mating and flight. This floral association is widespread in the genus, with species such as Cyclocephala ruficollis observed feeding on sorghum (Sorghum sp.) pollen in Colombia and on Citrus flowers in similar contexts. Other examples include Cyclocephala celata pollinating Caladium bicolor while feeding on its floral rewards in Brazilian rainforests, and Cyclocephala colasi interacting with thermogenic inflorescences of Philodendron solimoesense. In controlled rearing, adults of congeneric species are provided with fruits like apples or bananas, or native plants from collection sites, to sustain them, though field observations for some species like C. barrerai indicate limited feeding focused more on reproductive activities.^[24] Feeding in both life stages is influenced by environmental factors, such as soil moisture for larvae, which enhances root access and growth rates, and floral scents or heat for adults, which guide them to host plants at night. While larval root consumption can lead to plant wilting and turf damage, adult feeding rarely causes noticeable harm to ornamental or agricultural plants, emphasizing their role as incidental floral consumers rather than pests in aboveground contexts.^[22][27]

Pollination Interactions

Cyclocephala beetles, belonging to the tribe Cyclocephalini within the Scarabaeidae family, play a significant role in cantharophilous pollination systems, particularly for night-blooming flowers of early-diverging angiosperms. These beetles are primary pollinators for plants in families such as Annonaceae, Magnoliaceae, and Araceae, where they are attracted to floral scents, thermogenic heat, and rewards like petal tissues or pollen. Across nearly 500 cyclocephaline species, including over 300 in the genus Cyclocephala, floral associations have been documented with 58 plant genera spanning 17 families and 15 orders, predominantly in the Neotropics. Mutualistic interactions often involve beetles entering floral chambers at dusk, feeding on adaxial petal surfaces, and inadvertently transferring pollen between flowers while seeking mating or feeding sites.^[28] In Annonaceae species like Annona coriacea, Cyclocephala species such as C. atricapilla and C. undata serve as principal pollinators during nocturnal anthesis, which lasts 52–56 hours and peaks between 22:00 and 03:00. Beetles arrive via strong fruit-like scents, consume petal tissues within the floral chamber, and deposit pollen on stigmas, facilitating both cross- and self-pollination (geitonogamy) despite the plant's self-compatibility. Similarly, in Magnolia tamaulipana (Magnoliaceae), C. caelestis accounts for 52% of floral visits, feeding on carbohydrate-rich petals in thermogenic flowers that generate excess heat (up to 9.3°C in the female phase) to enhance scent dispersal and attract beetles. These interactions result in moderate pollination success, with only 44% of open-pollinated stigmas receiving pollen in M. tamaulipana, underscoring the beetles' efficiency in specialized systems.^[29][30] Beyond basal angiosperms, Cyclocephala paraguayensis pollinates Lagenaria siceraria (Cucurbitaceae) flowers, drawn by the sesquiterpene (3S,6E)-nerolidol, which comprises 97% of the floral volatile emissions (~1.3 μg/hour). Beetles aggregate at twilight for mating and pollen feeding, with 80% of females and 50% of males carrying pollen in their guts, confirming their role in pollen transfer. In Araceae like Philodendron acutatum, a single Cyclocephala species exclusively pollinates self-incompatible inflorescences, achieving over 90% fruit set through precise nocturnal visits. These examples highlight the beetles' adaptability in diverse pollination mutualisms, often enhanced by floral thermogenesis and specific attractants.^[31]

Predators and Parasites

Cyclocephala species, like other scarab beetles, face predation across their life stages, with eggs and larvae being particularly vulnerable in soil environments. Egg predation is significant in turfgrass settings, where up to 73% of Cyclocephala lurida eggs can be consumed within 72 hours of oviposition. Ants dominate this predation, with Solenopsis molesta accounting for up to 83% of attacks, including 54–68% in specific field studies, while other ants like Lasius spp. contribute 5–13%. Ground beetles such as Scarites subterraneus and rove beetles (Staphylinidae) also prey on eggs but at lower rates, typically less than 5%.^[32] Larval stages, known as white grubs, are targeted by soil-dwelling predators including ground beetles (Carabidae) and ants, which feed on the root-feeding grubs in turf and agricultural soils. Birds such as scarlet ibis (Eudocimus ruber), white ibis (Eudocimus albus), and white-faced ibis (Plegadis chihi) consume adult and larval Cyclocephala, incorporating them into wetland and grassland diets. For adults, which emerge to feed nocturnally on foliage and flowers, primary predators include birds, with additional predation from spiders, predatory insects, and small mammals in open habitats.^[33][11][34] Parasitism is a key regulatory factor, particularly for larvae. Hymenopteran parasitoids in the family Tiphiidae, such as Tiphia pygidialis and Tiphia vernalis, target third-instar grubs of species like Cyclocephala lurida and Cyclocephala signaticollis, with parasitism rates reaching 14–33% in turfgrass sites. These wasps locate grubs by antennal tapping, oviposit externally, and their larvae develop ectoparasitically, leading to host death. Scoliidae wasps also parasitize white grubs of Cyclocephala, though less frequently documented, serving as natural enemies in soil ecosystems.^[35][36][33] Nematodes are prominent endoparasites, with species in Thelastomatidae (e.g., new taxa from Cyclocephala signaticollis) and Rhabditidae (e.g., Parasitorhabditis platidontus and Cruznema campestris) infecting larvae in South American populations. Entomopathogenic nematodes like those in Steinernematidae interact with third-instar C. lurida grubs under controlled conditions, reducing survival. Bacterial pathogens, including Bacillus popilliae, infect Cyclocephala parallela larvae, causing milky disease. Fungi such as Metarhizium brunneum can infect larvae, demonstrating potential for biological control. Phoretic mites (Acaridae and Macrochelidae) associate with adults, potentially vectoring pathogens but primarily non-lethal.^[11][37][38]

Economic Significance

As Pests

Species of Cyclocephala, commonly known as masked chafers, are recognized as significant agricultural and horticultural pests primarily due to their larval stage, known as white grubs, which cause substantial root damage to various crops.^[18] These grubs feed voraciously on plant roots, leading to weakened plants that exhibit symptoms of drought stress, wilting, and eventual death if infestations are severe.^[39] Adult beetles, while less damaging, may occasionally feed on foliage or flowers, but their impact is generally minor compared to the subterranean larval activity.^[19] The primary economic concern arises from damage to turfgrass, where larvae sever roots, resulting in irregular brown patches and turf that can be easily rolled back like a carpet.^[40] In turf settings such as lawns, golf courses, and athletic fields, economic thresholds for intervention are typically set at 6–10 grubs per 0.1 m², beyond which visible damage and secondary issues like vertebrate predation exacerbate losses.^[18] For instance, in Kentucky bluegrass, damage becomes evident at 13–20 grubs per 0.1 m², highlighting the tolerance limits of well-maintained turf.^[18] In Florida, species like C. lurida and C. parallela produce two generations annually, with the first-generation grubs causing the most severe root feeding from July to August, leading to thinning and spongy turf.^[41] Beyond turf, Cyclocephala grubs infest row crops and horticultural plants, feeding on roots of corn, sorghum, sugarcane, and forage grasses, which can reduce yields and require costly management.^[19] In Neotropical regions like Brazil, species such as C. distincta damage sugarcane roots, contributing to yield losses in major production areas.^[42] In blueberry plantations, particularly affecting young plants, larval root pruning induces drought-like symptoms and plant mortality, necessitating integrated pest management to mitigate establishment risks in new orchards.^[39] These pests are widespread across North America, with species such as C. borealis prevalent in the Midwest and Northeast, C. lurida in the South and Midwest, and C. parallela in Florida, contributing to regional agricultural losses through direct crop damage and control expenses.^[18]

Ecological Roles

Cyclocephala beetles, commonly known as masked chafers, play significant roles in tropical and subtropical ecosystems, primarily through pollination by adults and soil health contributions by larvae.^[43] These scarab beetles (Coleoptera: Scarabaeidae, Dynastinae) are integral to the reproduction of early-diverging angiosperms, where their foraging behaviors facilitate cross-pollination.^[28] Adults of numerous Cyclocephala species serve as primary pollinators for plants in families such as Annonaceae, Araceae, Arecaceae, Cyclanthaceae, Magnoliaceae, and Nymphaeaceae, among others.^[28] For instance, Cyclocephala colasi is a key pollinator of Philodendron solimoesense (Araceae), where beetles are attracted to thermogenic inflorescences that provide warmth, shelter, and food rewards like pollen and floral tissues.^[44] Similarly, species like Cyclocephala lurida pollinate Dieffenbachia species (Araceae) and contribute to the reproductive success of Nymphaea (Nymphaeaceae) by transferring pollen while feeding on nectar and floral parts. These interactions often involve specialized floral scents, such as (3S,6E)-nerolidol, which act as attractants, underscoring the beetles' role in maintaining biodiversity in Neotropical forests.^[31] At least 97 Cyclocephala species participate in such mutualisms, supporting pollination across 58 plant genera in 17 families and 15 orders, predominantly in the New World.^[28] Larvae of Cyclocephala species contribute to belowground ecosystem processes by feeding on roots, organic matter, and decaying plant material, thereby aiding decomposition and nutrient cycling.^[43] In agricultural and natural soils, species like Cyclocephala lunulata exhibit saprophagous habits, breaking down leaf litter and enhancing soil fertility through the release of nutrients such as nitrogen and carbon.^[43] This activity promotes bioturbation, improving soil porosity, structure, and aeration, which facilitates water infiltration and microbial activity essential for ecosystem productivity.^[43] For example, Cyclocephala barrerai larvae process organic detritus, recycling nutrients back into the soil and supporting plant growth in tropical environments.^[26] Overall, these larval functions help sustain soil health, indirectly benefiting primary productivity and carbon sequestration in diverse habitats.^[43]